Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. xHo. oe.adpyoeeial sltdfo h type the of from This isolated species prep.). phylogenetically and in Copel. Hook.) al. genus species) ex (four et small the Windham to sister 2009; is hemionitid group al. the et within (Windham lineage that species ferns diverging twenty about early of clade an a represents 2011), al. et Li to 2004; Smith relationships “ the called whose been has clades genera supported cheilanthoid Cheilanthes well other among diverse, or to 2011). 1998; within al. nested et anomalous 1995, Eiserhardt morphologically species single, 2008; 2007; from al. Kirkpatrick gamut Rollo the et 2007; run Rothfels These al. 2007; and et al. Zhang et (Gastony Prado 2007; al. ferns et Schuettpelz cheilanthoid clade major every in of appearing genus this genus to assigned tionally defined of loosely polyphyly extensive and the large the Cheilanthes is of ferns boundaries cheilanthoid the 2008; 2011), al. al. et et unchanged. Link- remain Eiserhardt genera 2010; Rothfels cheilanthoid most 2011; Schneider 2007; al. and al. et Yesilyurt et Perez 2009; 2007; al. Zhang Kirkpatrick and et 2007; 2007; Windham (Gastony al. al. monophyletic et et not Schuettpelz Prado most 1998; are that 1995, genera indicating Rollo major studies the phylogenetic grow- of a of Despite number 248). generic 1982: ing natural Tryon and and practical contentious (Tryon a most to classification” Rollo “the respect with and as ferns to (Gastony of referred group group been has this which in 1998), rampant is a convergence (Pteridaceae), Morphological ferns. in by diversified avoided ferns have normally habitats that semi-arid species cheilanthoid + 400 of involves group monophyletic trend this to Sm.; Sundue; & Labiak Moran, Alansmia O 10.1600/036364412X656626 DOI © Botany Systematic re.Ti a eutdi h eonto fmn e genera new many of (e.g. ferns recognition polypod the the across in resulted has This towa aries. directed substan- been and 2012), has al. effort et tial Rothfels al. 2011; et al. et (Smith Christenhusz overhauled 2006; been have classifications mo of Family-level pace rapid the to part, large in oyih 02b h mrcnSceyo ln Taxonomists Plant of Society American the by 2012 Copyright h anipdmn ognrcrognzto of reorganization generic to impediment main The ensseaishscagdrdclyi h atdcd due, decade past the in radically changed has systematics Fern iueyblaemriso auelae v.sot t40 at smooth (vs. leaves mature of margins group. bullate the minutely within evolution reticulate nuclear: oohltcgopo pce ihntehmoii ld nomlycle h “ the called informally genus clade hemionitid (the the relatives within closest species of group monophyletic Cheilanthes eete e obntosaemd,adtonwspecies, new two and made, are combinations new seventeen prs(s 2salo 6lre e prnim n sal lbosla lds oa fnnte pce r eonzdwithin recognized are species nineteen of total A blades. leaf glabrous usually and sporangium, per large) 16 or small 32 (vs. spores from distinguished is genus Araiostegiella Abstract— Keywords— .Kslr oul ude&Labiak; & Sundue Moguel, Kessler, M. w eetpyoeei tde aedocumented have studies phylogenetic Recent Sw. .s r euu tbs.Aogtelte swhat is latter the Among best. at tenuous are s. s. Cheilanthes 21) 74:p.845–860 pp. 37(4): (2012), gapCp stenwydsrbdgenus, described newly the as non oeua hlgntcsuiso hiatodfrscnimta h genus the that confirm ferns cheilanthoid of studies phylogenetic molecular Ongoing oehrwt pr aat icmcietemrhlgcladgorpia onaiso h e eu n investigate and genus new the of boundaries geographical and morphological the circumscribe to data spore with together ) Gaga hiatodfrs yrdzto,rtclt evolution, reticulate hybridization, ferns, Cheilanthoid .Kt stui.Oentbeexception notable One Tsutsumi). & Kato M. hiate marginata Cheilanthes , 1 eateto ilg,Dk nvriy uhm ot aoia278 .S A. S. U. 27708, Carolina North Durham, University, Duke Biology, of Department .micropteris C. e enGnsSgeae from Segregated Genus Fern New a , Leucotrichum Cheilanthes Aspidotis Moranopteris a-e Li, Fay-Wei Cheilanthes drassiggnrcbound- generic reassessing rd eua hlgntcstudies. phylogenetic lecular n hlgntclydsatfo h yeseisof species type the from distant phylogenetically and ) .s yissrnl ifrnitd nrmria suonui,tepouto f6 ml r3 large 32 or small 64 of production the pseudoindusia, inframarginal differentiated, strongly its by s. s. Labiak; Gaga w Krptik2007; (Kirkpatrick Sw. .l,wt pce tradi- species with l., s. Gaga 2 uhrfrCrepnec ([email protected]) Correspondence for Author .Y ia .Prado; J. & Hirai Y. R. 1 ru”(ikland (Mickel group” sdsigihdfrom distinguished is ahenM Pryer, M. Kathleen nti td,w s oeua aafo orDArgos(plastid: regions DNA four from data molecular use we study, this In . omnctn dtr hisnGemmill Chrissen Editor: Communicating Serpocaulon Aspidotis Mickelia (Nutt. aagermanotta Gaga .C. R. .R. A. + ,adls rmnnl utosadsraeaailbaesrae.Tenew The surfaces. blade adaxial striate and lustrous prominently less and ), 845 Aspidotis matK 1,2 “ aafo he lsi ein ( regions plastid three from data n rps e pce ee aooyfrti group this for taxonomy genus level ferns. the species cheilanthoid within of new evolution a propose reticulate and of the patterns from sequences examine use also We mater- nuclear hybrids. identifying of in progenitors species assist nal estimate and group, relationships, the and of numbers monophyly the assess to used 21) aeil o N xrcinwr bandete from either obtained ( were Database loci three Lab for plastid Fern Sequences extraction (fernlab.biology.duke.edu). vouchered archive the tissue al. from DNA silica-dried et or Li sheets by for herbarium study accessioned barcoding DNA Materials a in 58 (2011). included study; previously phylogenetic were molecular the these for of sampled were isotypes) five and i ) oa f7 pcmn Apni )rpeetn l known all representing 1) (Appendix specimens 75 “ the of from total (Appen- species study A molecular the 1). for sampled dix also were which of 42 included, asstegenus the passes among “ the relationships Cheilanthes of phylogenetic inclusion ferns, of cur- cheilanthoid our Given understanding prep). in al. rent et Windham 2011; al. et Eiserhardt erdciemd fec pce ntegns Sequence genus. and the level in ploidy species extensive each the to of estimate we mode due to study, reproductive measurements this uncertain actual spore In the is apomixis. use and and involved 2011), polyploidy, al. taxa 2007; hybridization, et of (Kirkpatrick Eiserhardt 2008; studies number al. sam- phylogenetic et been has Rothfels previous one only in 2004), Smith pled and (Mickel group this the Treatment). segregate Taxonomic to chosen have “ we group, cohesive phologically Because “ genus. distinct the subsuming under include group” options disruptive less (W morphologically undefinable (e.g. genera well-characterized diverse, several subsume combinations, new of expansion an Such hiate marginata Cheilanthes .marginata C. eio oyliy pr measurements. spore polyploidy, Mexico, , ao Sampling— Taxon n ihe .Windham D. Michael and lhuhaottet pce aebe trbtdto attributed been have species twenty about Although yisruddt teut v.mcoae emn apices, segment mucronate) (vs. attenuate to rounded its by and gapCp .marginata C. aamonstraparva Gaga Cheilanthes matK sutnbebcueterslatcaeas encom- also clade resultant the because untenable is Doryopteris ru”a e eu eennamed herein genus new a as group” Aspidotis , ein ncnucinwt h pr aa to data, spore the with conjunction in region, hiate marginata Cheilanthes rbcL, Hemionitis Cheilanthes aeil n Methods and Materials o pr nlss oa f4 ape rmthe from samples 44 of total a analyses, spore For ru”i eesont edsic rmits from distinct be to shown here is group” and ru”adthree and group” Cheilanthes .S.and Sm. J. Aspidotis Hemionitis issse ld)o eonzn ta a as it recognizing or clade) sister (its trnG-R hsgopi eesgeae from segregated here is group This . r described. are , hc a roiyover priority has which , (Pteridaceae) n n ula ou ( locus nuclear one and ) Peiaee splpyei.A polyphyletic. is (Pteridaceae) sarltvl itntadmor- and distinct relatively a is nhme l npe..Other prep.). in al. et indham 1 Adiantopsis matK ol eur udesof hundreds require would ru”(nldn n holotype one (including group” matK .marginata C. Aspidotis , rbcL , rbcL Fe and , ´ ) n ol be would and e), , trnG-R pcmn were specimens .marginata C. trnG-R Cheilanthes gapCp ru”in group” Gaga Gaga ;and were ) ; are ) (see . Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. pcmnicue ntesuy yoeei nlsso w species two of fertile analyses each Cytogenetic of mode study. in reproductive included the now the in infer included to specimen sporangium per spores eedsesdi lcrldo nasie pr mgswr then were images Spore slide. a Beck on sporangia 80 drop intact 2009; nearly glycerol at al. or intact a captured et from in then Spores Grusz 2011). dispersed and al. smallest 1993; were et triploids, Sigel the Rabe 2010; apomictic have and al. et diploids (Windham tetraploids, level; polyploids sexual ploidy higher by demon- with followed been correlated has spores, size be spore to ferns, cheilanthoid strated In specimen. fertile each num- mode. deter- spore previous reproductive with of estimated congruent minations sporangium), groups lost the non-overlapping or two open Although destroyed formed breaking are bers microscope. of usually spores process dissecting (some the 125 64 in or MZ 32 sporangium per exactly Leica spores totaling a of rarely bro- number using and The needle. slide estimated a microscope of was a tip the on were using glycerol open These of ken specimen. droplets each individual from in removed placed were infer sporangia one To analyses, intact data). molecular three unpubl. the to in Windham included D. specimens (M. for produce sporangium mode reproductive per apomicts spores whereas 16 sporangium or per 32 spores a 64 produce of individuals preponderance reproducing sexually ferns, leptosporangiate most of species two locus, nuclear 4 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC including analyses, plastid eight 2011) the al. for et selected Eiserhardt micropteris 2009; were C. al. species et ferns outgroup cheilanthoid Windham of 2007; studies phylogenetic (Kirkpatrick previous on Based generated. 846 hoooecut sdt airt liylvldtriain.Tenmesascae ihseisnmsaeteFr a aaaeaccession Database Lab Fern clade). for kaulfussii the specimens = are K voucher names clade, identify purpusii species major arrowheads = with the P Grey identify clade, associated arrowheads. bars marginata numbers error = in the The M terminating below clade, determinations. letters lines angustifolia capital level fine the ploidy and by calibrate (fernlab.biology.duke.edu), connected numbers to are used individual counts single chromosome a by produced sporangia ln hi ogs xst eemn h vrg pr eghprspeci- per length spore of average measured specimens the were determine Two specimens) to men. apomictic axis longest (for their specimens) sporangium sexual along per (for spores sporangium per 10 ImageJ spores or Using 20 microscope. least dissecting at 125 2011), MZ (Rasband Leica a on mounted camera erdcieMd n liyAssessment— Ploidy and Mode Reproductive eas eemndaeaesoesz oass h liylvlof level ploidy the assess to size spore average determined also We Fig. .Rneo vrg pr egh ( lengths spore average of Range 1. tetp pce of species type (the + Gaga anfcto sn ao O ee S digital XSi Rebel EOS Canon a using magnification ( .arizonica G. Gaga Aspidotis n he of three and and Cheilanthes eeicue soutgroups. as included were .cuneata G. Aspidotis m )i pce of species in m) ;frteaayi,o the of analysis, the for ); eue h ubrof number the used We niaeta,a with as that, indicate ) Apni ;Fg 1) Fig. 1; (Appendix Gaga and Aspidotis egho 0–5 aepis.Bt mlcn eeotie rm77% from total obtained a were of the amplicons in (out Both specimens overlap the pairs). two of of base the pairs 600–650 and base of 2008), a 400 length al. yielded approximately two et both had yield (Schuettpelz amplicons combinations band to copy-specific primer 2008), “short” new al. single The et amplicons. (Schuettpelz overlapping ESGAP11R1 and ESGAPCP8F1 e nenlpies WHaF (5 FWCHgapF1 primers, internal new hte etwscnutduigR( eeomn oeTeam Core Mann- Development size. A (R spore in size. R differences chromosome observed spore using of and significance with conducted the data test level was to length ploidy 2008) test spore correlate U average to Whitney used both were had counts we which for a eie rma5%mjrt-uecness nterr vn when event rare clade the In that consensus. consensus of majority-rule The sequences 50% a constituent allele. from the one derived was represented represent least then the to at that On with considered sequence clade 2002). was recognizable (Swofford and sequences v4.0a123 minimum two each PAUP* tree, in MP out unrooted the (MP) carried parsimony to maximum was A according (2009). analysis sequences al. et colony Grusz by these established protocol from determined were each Alleles 1). Appendix 12; from mean: 7–18, from (ranging specimen sequences each for obtained alignment were resulting single a (Mu into v0.9971 The aligned PhyDE manually using primers. and pooled M13R then done were specimen were and reactions M13F sequencing using subsequent Wisconsin). and (Madison, amplifications kit cloning Colony pGEM-T Promega the using separately 3 TTTTG-3 eke l 21)wr sdwt w diinlsqecn primers, sequencing additional two with used (5 were (2010) trnGIF3 al. et Beck plastid for 2007). (2011); plastid Pryer of and sequencing (Valencia, (Schuettpelz and kit Amplification protocols mini published plant DNeasy following QIAGEN California) the using extracted was DNA AATTCGTTCCTTATT-3 0 ,wr eindt epcieypi ihtepbihdprimers published the with pair respectively to designed were ), N xrcin mlfcto,Coig n Sequencing— and Cloning, Amplification, Extraction, DNA iherrbr niaigoesadr eito.6-ad32-spored and 64- deviation. standard one indicating bars error with , 0 n WHaR (5 FWCHgapR1 and ) 0 -AATAAGGAACGAATTAARG-3 trnG-R Gaga gapCp 0 ¨ ld ihnwihteseie slctd( = (A located is specimen the which within clade .T mlf h nuclear the amplify To ). h rtcl fNglnu ta.(07 and (2007) al. et Nagalingum of protocols the , lre l 00.A es ee ooysequences colony seven least At 2010). al. et ller aae.Each dataset. 0 -CTYCCAGAGCTCAATGGTAAACT- 0 -TTGCTAAGGGCAATGCCTGC matK gapCp 0 n rGR (5 trnGIR3 and ) and rbcL mlcnwscloned was amplicon gapCp olwdL tal. et Li followed ein two region, Genomic 0 -CYTT Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. angustifolia halliae 46.54 of length 4soe.Atog ili hoooecut aenot from have reported counts been chromosome yet diploid Although 20 64-spored. includes 1) Fig. of side (left cheilanthoid sporangium most groups sharply-defined ( per among two form spores measurements taxa Spore large ferns. apomictic 32 of the characteristic only Twenty containing produced spores. sporangia mens unreduced 32-spored presumably rare, produced larger, also 1) Fig. ( latter the and of produc- 6550 Two sporangia. their 64-spored on of based tion reproducing sexually as categorized of specimens Forty-three isn aa()00 .725 .613.27 27.78 G + TIM 781 781 0.96 21.72 Mixed 3,747 3,890 28.08 G + 2.52 GTR 1,136 1,279 G + I + GTR 9.24 0.47 1,309 1,309 G + TVM 28.73 0.08 1,302 1,302 ln Best used Model (%) sites Variable (%) data Missing included Characters length Alignment 847 topological the among no observed was Because 70) to > support. rbcL, BS set branch threshold a “genthreshfortopoterm” determine (using with incongruence done to with was replicates (BS) 2006) Bootstrapping 1,000 out bootstrapping). carried (Zwickl for were (100,000 v2.0 analyses 1,000,000 second-best-fit FERNS CHEILANTHOID (ML) the Garli OF likelihood 2001), GENUS using Maximum NEW Ronquist used. A and available GAGA, was not (Huelsenbeck AL.: the model was ET v3.2 under model LI 2002) MrBayes best-fit the Swofford in When 1998; criterion. Crandall information and Akaike (Posada ( Modeltest evolution dataset using Sequence each analyses. phylogenetic for to models prior excluded were these the only manually; sequence. consensus the a in making identified to prior easily were removed and these alignment sequences, colony chimeric were there 2012] aae htwste ujce oM n aeinifrne(I.For (BI). inference Bayesian ( and dataset ML plastid to combined subjected the then was that dataset liylvl All level. ploidy Aspidotis nysihl agrta hs fkondpod ntesister the in diploids known of those than larger slightly only aae.Tecmlt euneainet r eoie nTreeBASE in plastid deposited the in with are combined TB2:S12433). alignments assist number be (study sequence not to could complete it present The taxa, serves were dataset. polyploid the dataset alleles of multiple most nuclear phylo- and for maternal contributions the overall biparental Because the determining con- provide were framework. to datasets plastid combined genetic The therefore partition. and single a gruent as (PP). but probabilities above posterior described discarded calculate output was to nuclear sample Drummond used the the was the and For rest of runs, the 25% (Rambaut and MCMC first burn-in v1.5 The the as convergence. Tracer After ensure in to partitions. 2009) inspected the were among was parameters prior vary substitution rate the to dataset, and partitions plastid set loci the sampled combined across were unlinked the trees were For parameters and station- generations. rates, the substitution 1,000 both the for every and distribution Dirichlet frequencies flat state (one a ary followed chains with Priors settings four generations. million default with 13 the for each running Two was out, cold) three 2001). BI carried and heated Ronquist locus. were and runs each (Huelsenbeck MCMC to independent v3.2 assigned MrBayes model using best-fit performed the with partitions, three p hlgntcAnalyses— Phylogenetic erdcieMd n liyAssmn Fg 1)— (Fig. Assessment Ploidy and Mode Reproductive Table < .01i anWinyUts;Fg ) n ihamean a with one 1), Fig. test; U Mann-Whitney in 0.0001 and L i.1,adteohrwt 66.26 with other the and 1), Fig. ; .Sqec hrceitc,bs-i euneeouinmdl,adrsligte ttsis isn aade o nld indels. include not does data Missing statistics. tree resulting and models, evolution sequence best-fit characteristics, Sequence 1. trnG-R eeaaye oass erdciemd and mode reproductive assess to analyzed were .apiacea G. 50to 6550 gapCp aaes h he ein eecmie noone into combined were regions three the datasets, trnG-R m Aspidotis (from m aae,M n Iaaye eecridotas out carried were analyses BI and ML dataset, .hirsuta G. 86 dniidb ogbakarw in arrows black long by identified 7876; matK aae a miuu lgmn ein and regions alignment ambiguous has dataset o ahdtst l eune eealigned were sequences all dataset, each For Gaga matK n 1o the of 21 and Gaga Results , sioi californica Aspidotis rbcL 48228 29164 42781 1,5.17–3,074.8286 –12,156.1157 –4,297.8015 –2,981.6344 –4,802.2783 plastid pr ie nti ru are group this in sizes spore , 13.Tesalsoe group small-spored The 5133). Gaga + , n he ftesse genus sister the of three and rbcL trnG-R matK ape l fwihwere which of all samples + ,and trnG-R Gaga gapCp ,M a u with run was ML ), m pcmn were specimens (from m to .angustifolia G. eechosen were ) Gaga .carlotta- A. plastid speci- rbcL Gaga matK , h aasgetta h atrmyb oecoeyrltdto related 3). 2, closely (Figs. more clade marginata/purpusii be the may to than latter other the each (i.e. that suggest groups though data monophyletic unsupported, between the are two clades) Relationships kaulfussii other and the angustifolia clades. the and clade purpusii larger and this marginata the ( prnim(i.1.Bsdo h vial aa ehypoth- per of we plant spores data, this available that large the esize on 64 Based producing 1). (Fig. in sporangium examined specimens the Fg.2 ) ohtepatd( plastid the Both within purpusii, 3). resolved 2, marginata, are (Figs. clades, the kaulfussii designated and angustifolia, here groups, sites phyletic variable many non-coding as the with marker as useful particularly sister a proves be its 2011), al. to is et (Kuo toolkit as phylogenetics fern the analyses, to added 3) (Fig. nuclear to relationship and 2) (Fig. plastid “ etfteouinmdl,adte ttsisapasin appears statistics genus new tree the of and monophyly The characteristics, 1. models, Table sequence of evolution summary a best-fit study; phylogenet- this analyzed for were ically nuclear) one and plastid (four ces 2soe e prnimaeaigmr hn60.5 than more averaging sporangium all per 1), spores 32 Fig. ( of side one (right group but large-spored the to belonging poal l)o hs ape ersn euldpodspo- diploid spores. tetraploid sexual haploid producing represent documented rophytes samples these the of all) have of (probably group those this carlotta-halliae of genera than A. members related All smaller closely prep.). spores other in al. in et diploids (Windham sexual known for genus infcnl agrta hs ftetraploid of those than larger significantly harrisii ( G. distribution in large-spored two 1) the The of Fig. end range. in size arrowheads this grey known span of by specimens (indicated voucher triploids the because apomicts triploid anomalous the (excluding oeitrsig hssml of sample This interesting. more diploid sexual the of diploid end sexual to low size the in Because (similar at range 1). spores Fig. produces in (indicated also arrow sporangia black 64-spored that long plant of by a predominance by a produced sporangium showed 32-spored rare 1). a (Fig. from triploids from sample apomictic spore known particular of This those than smaller yet erpod,wihmkstedt for be data sexual the should by makes produced they spores which diploid such, tetraploids, the to As repre- size spores. in sporangium comparable 32-spored diploid this unreduced, of sent contents the that esize gapCp hiate marginata Cheilanthes hlgntcAnalyses— Phylogenetic mn h 23 the Among plastid aaessrnl upr itrrltosi between relationship sister a support strongly datasets ) Aspidotis .harrisii G. 54 eev pca eto.Bt aespores have Both mention. special deserve 5574) trnG-R Aspidotis Fg )adwl ihnterneobserved range the within well and 1) (Fig. Gaga Fg ) n ehpteieta most that hypothesize we and 1), (Fig. trnG 54 r 2soe.Seieswith Specimens 32-spored. are 5574) .harrisii G. ru” srbsl upre nboth in supported robustly is group”) - ape ihkonsoenumbers spore known with samples h plastid The . R ein(al ) ormjrmono- major Four 1). (Table region .apiacea G. lsi cmie)nuclear (combined) plastid ieDAainetdt matri- data alignment DNA Five matK Gaga sasxa tetraploid. sexual a is + .harrisii G. .angustifolia G. rbcL matK pcmn ttelower the at specimens as interpreted are 7876) .angustifolia G. Aspidotis + .harrisii G. trnG-R .angustifolia G. ee nyrecently only gene, Gaga suiu among unique is .carlotta-halliae A. ,w hypoth- we ), frel the (formerly )andnuclear 50derives 6550 54even 5574 50and 6550 m long m Gaga Gaga 6550 gapCp , Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. aeinpseirpoaiiy + qas10 r10.Nmesascae ihseisnmsaeFr a aaaeacsinnmes(fernlab. numbers accession Database Lab sexual = Fern inferred indicate Ar are circles Abbreviations: open names and respectively. species = grey, triploid, An Black, with apomictic “Type.” associated superscript and the Numbers tetraploid by 1.00. sexual identified are or specimens 100% type nomenclatural equals biology.duke.edu); “+” probability; posterior Bayesian 4 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 848 Fig. .angustifolia G. .Tebs Lte rmteaayi fteplastid the of analysis the from tree ML best The 2. 51 o= Mo 6541, .monstraparva G. . matK + rbcL + aaarizonica Gaga trnG - R aae.Bac upr ssona ahnd sM bootstrap/ ML as node each at shown is support Branch dataset. ,Me= .membranacea G. ,G= .germanotta G. ,D= .decomposita G. diploid, , Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 02 IE L:GG,ANWGNSO HIATODFRS849 FERNS CHEILANTHOID OF GENUS NEW A GAGA, AL.: ET LI 2012] ubrrpeetdfeetallsrcvrdb cloning. by recovered alleles different represent number Fig. .Tebs Lte rmteaayi ftenuclear the of analysis the from tree ML best The 3. gapCp aae.Smosa nFg .Triassaigtesm ao aeadaccession and name taxon same the sharing Terminals 2. Fig. in as Symbols dataset. Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. slkl,ti eainhpi oryspotd(8 MLBS/ (78% supported clade 2). poorly Fig. angustifolia PP; is the 0.76 relationship to sister this as likely, clade is kaulfussii posi- the species, phylogenetic of the single tion Although a sampled. specimens of five consists all clade kaulfussii kaulfussii The G. species, new 2). a Fig. to in sister hybrid is that probable group the chaerophylla and G. ( monophyletic, them as between resolved are + ld,awl-upre al ihtm separates dichotomy angustifolia early the Within well-supported species. angustifolia three a other the clade, to sister as and pellaeopsis G. srsle smnpyei ntepatddtst Both dataset. plastid the in pellaeopsis monophyletic G. as resolved sister is turn, in are, species two these to and similarity, logical harrisii species. this G. var. within taxa as infraspecific identified of collections among nested is complanata 2) Fig. in 8103 stp of isotype membranacea G. with 5 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 850 sa sebaeo pmci lorpod ihdiverse with allotriploids apomictic of “species” assemblage this that an indicating 3), is (Fig. alleles distinctive through of two, samples originated of nine isotype that an all (including triploids contrast, apomictic By autopolyploidy. suggesting are sporangium, they per spores that homozy- large apparent 32 these produced of gotes Two 3). (Fig. sampled specimens uinta htosre ntepatdpyoey nyone Only phylogeny. plastid the gapCp in observed that than lution the 07sae identical shares 8097 membranacea G. across identical nearly are sequences plastid inosre n hi mlctosfrtxnm r elabo- Discussion. are the taxonomy in for rated implications evolu- their reticulate and we of observed patterns Here tion the datasets; the event. nuclear from and inferred hybridization plastid results a phylogenetic the in present The separately involved about progenitors. information parents reveal maternal can both hand, in other identifying the inherited on for data, plastid nuclear maternally useful 1992), are be Yatskievych and data to Because (Gastony genus. shown ferns the cheilanthoid been in have inves- evolution to plastids data reticulate nuclear and potential plastid 2004). tigate of Smith combination and a used (Mickel We clades angustifolia and marginata nFg ;seTxnmcTetet,i iial divergent minimally is Treatment), Taxonomic from see distinct. 2; phylogenetically Fig. be in to here germanotta shown hand, Gaga of is synonym other 2004), a Smith the as treated On recently differences. phological hirsuta G. .monstraparva G. ihnteprui ld,neither clade, purpusii the Within N P the both in common be to reported been has Hybridization .decurrens G. LASTID gapCp UCLEAR .purpusii G. .germanotta G. .membranacea G. leewsrcvrdi aho h five the of each in recovered was allele and , aapoiesgiiatymr hlgntcreso- phylogenetic more significantly provide data hiate complanata Cheilanthes P r lsl eae,cnitn ihtermorpho- their with consistent related, closely are n hspoie ospotfrterecognition the for support no provides thus and + P HYLOGENY n t lsi eune r dnia across identical are sequences plastid its and , omn oyoywt cesosof accessions with polytomy a forming HYLOGENY .chaerophylla G. All . . and .arizonica G. u ihadfeetptra rgntr The progenitor. paternal different a with but e yrdseisdsrbdhr (“G” here described species hybrid new a , sismtra rgntr Because progenitor. maternal its is .decomposita G. and , Neither . .cuneata G. .hintoniorum G. ttomyb yrdinvolving hybrid a be may too it , (F (F matK IG .hintoniorum G. “e nFg ) niaigthat indicating 2), Fig. in (“Me” hiate spiculata Cheilanthes IG )Wti h agnt clade, marginata the 2)—Within . )Wti h agnt clade, marginata the 3)—Within . ept hi osdrbemor- considerable their despite , .angustifolia G. + , pcmn omamonophyletic a form specimens .decomposita G. D nFg )gop with groups 2) Fig. in “D” ; rbcL, var. .arizonica G. r oohltc with monophyletic, are .apiacea G. interrupta and .monstraparva G. aacomplanata Gaga togysupported strongly nor trnG-R hwda least at showed ) from nor .chaerophylla G. ( .marginata G. .complanata G. .marginata G. Mce and (Mickel .lerstenii G. .purpusii G. sequences .cuneata G. .hirsuta G. .apiacea G. Gaga (“Mo” and sp. G. , , ei oiino h alusiclade. kaulfussii the of position phyloge- netic unresolved the regarding information additional no ik ebro hsgop(is ,3.Crsesn(1906) Christensen 3). 2, (Figs. group this treated generic of published member invalidly a Link, An past. the name, in to alluded was snwgnr.Hr efcso n ftelre potential larger the of “ the one as on known focus informally segregated we segregates, or Here genera genera. other new to transferred as be as to recognized need will groups past species the of many hoet ra l 0 hiatodfrsa igehighly single name a the we as bear ferns Unless would cheilanthoid (which 2011). al. genus + al. et polymorphic 400 et all Zhang Eiserhardt treat 2007; to 2008; al. choose al. et et of Schuettpelz Rothfels 2007; clade 2007; al. major et every Kirkpatrick Prado 1998; 2007; in 1995, Rollo appearing and (Gastony 1982). genus ferns cheilanthoid this Tryon to assigned and extensive of the Tryon documented polyphyly have studies 1981; phylogenetic changing Recent (Stolze constantly its circumscription and distinctiveness morphological notogop “r nFg ) n omn monophy- in a observed forming occasionally one to allele 3); unique Fig. group in letic (“Ar” fall that groups alleles two have species into this of samples Both allotriploid. obntoso rgntrgnms h morphologically The genomes. distinctive progenitor of combinations nteagsioi ld.Tenuclear The groups clade. well-supported angustifolia the the of in from any with divergent unaffiliated slightly other one the 3), Fig. in (“Mo” origin. hybrid a suggesting with both group which that of two to other resolved, clade, are to angustifolia groups correspond the supported Within highly phylogeny. consistent four clade, plastid the in the species other with all to sister as supported pellaeopsis G. of specimen only to related The was tetraploid. of sexual sample a complanata be G. may cies harrisii G. Two yosweesohr rwallsfo ohgroups. both from alleles homo- of reciprocally draw relationships be others Phylogenetic to appear whereas 2). specimens (Fig. zygous some dataset 3); plastid (Fig. the by contrast purpusii presented in Gaga 3), polytomy (Fig. the monophyletic to as resolved robustly each i.3,oeo hc ssse oalmmeso h marginata the in with of grouping members (“Me” all types to sister allele is clade. distinct which of two one have 3), Fig. and are another of clade one samples marginata to two the of The members unclear. other its to though monophyletic relationships as supported strongly is diploid, ual synonymized). been has Cheilanthes h eaiedsicieeso h “ the of distinctiveness relative The ihnteprui clade, purpusii the Within gapCp aagermanotta Gaga Othonoloma Othonoloma .cuneata G. ogun ihtehptei htti spe- this that hypothesis the with congruent , .lerstenii G. .arizonica G. lee eercvrdfo u nysml of sample only our from recovered were alleles eanucran although uncertain, remain ile single a yielded lee alit w ihyspotdgroups supported highly two into fall alleles Peiaee sntrosfrislc of lack its for notorious is (Pteridaceae) .arizonica G. .angustifolia G. .membranacea G. Cheilanthes . a ple yLn 13)to (1833) Link by applied was , aadecomposita Gaga sasnnmof synonym a as a stp)hdto ete fwhich of neither two, had isotype) (an .angustifolia G. .arizonica G. a he lee “”i i.3,two 3), Fig. in (“G” alleles three has Discussion aadecurrens Gaga loapast ea apomictic an be to appears also udrwihti pce recently species this which (under .l,wt pce traditionally species with l., s. aamonstraparva Gaga n to one , .purpusii G. n n with one and gapCp .membranacea G. .aica .purpusii, G. apiacea, G. .marginata G. h te dnia oan to identical other the , Cheilanthes “”i i.3 a alleles has 3) Fig. in (“D” n with and .chaerophylla G. leeweesteonly the whereas allele .marginata C. gapCp nerdt easex- a be to inferred , .marginata C. and .hintoniorum G. Cheilanthes aae provides dataset .marginata. G. and .chaerophylla G. a w alleles two has .cuneata G. soitn it associating , .apiacea G. r identical are Hemionitis group.” .hirsuta G. .hirsuta C. n the and , group” nthe in and and are is ), . , Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 02 IE L:GG,ANWGNSO HIATODFRS851 FERNS CHEILANTHOID OF GENUS NEW A GAGA, AL.: ET LI 2012] ifrn rmtoeosre ntetp pcmnof specimen type the on observed are hirsuta those G. surfaces pebbly from with different 4A) taxonomy. (Fig. pseudoindusia species from spiculate distinct pre-2004 3), and 2, the cohesive (Fig. be exist, to data intermediates molecular return some the Although a of advocate light in we names four these spiculata C. of treated nym (2004) Smith Beitel and complex, Mickel and whereas this spe- two Mickel in accepted (1995) (1981), cies Moran and Smith Yatskievych and conflicts. (1988), and inevita- taxonomic apomixis, to leading ble hybridization, 2004), Smith and and by (Mickel polyploidization obscured deltate, mostly seriously have are to variants appear broadly been boundaries These species and 4A-B). however, to or (Fig. continuous, spiculate pebbly to ovate slightly hairy from only leaf vary from with that margins significant, pseudoindusial is ranging complex this blades in observed diversity ihtespecies the with ugns epciey of respectively, used subgenus, later (1979) Reeves and icmcie h “ the circumscribed Kunth, pce ntegns nldn w aanwt science to new taxa nineteen two ( synonymy of including genus, total ( the a in recognize genus, species We and described newly prep) Treatment). a in as Taxonomic it al. clade segregate et to sister chosen Windham its from 2011; distinguished of al. easily species et Eiserhardt type 2007; the from isolated “ the monophyly of the demonstrate robustly reconstructions only 2011;genetic are al. two et last prep). Eiserhardt the in al. and 2007; et Windham lineage, (Kirkpatrick [= this related fifth to distantly the sister 3), is 2, Lellinger] (Figs. related another closely are one four first to the Tryon, and Tryon by mentioned and kaulfussii C. erdciemd n liylvlso la phyloge- clear a in show clade. pattern level kaulfussii netic are ploidy the clades and of major mode position the Reproductive kaulfussii these for among except and relationships resolved, well the angustifolia, 3); 2, supported purpusii, (Figs. highly marginata, four with clades: topologies congruent revealed Fg AB ,idctdb lc rohas;teexceptions the rachises arrowheads); include or black here by costae indicated and the D, 4A-B, along decur- kaulfussii, (Fig. extend long have that angustifolia, also pseudoindusia clade this the rent in taxa in Most bicol- clades. inconspicuously observed purpusii or scales concolored the ored with These scales from margins. brown rhizome differ conspicuous bicolored scales and have stripe 3) central dark 2, a (Figs. clade this to clades. these of each for below discussed are taxonomy hybridization, and mode, 1–3). reproductive (Figs. polyploidy, clades Morphology, kaulfussii and in dominate angustifolia, triploids marginata, apomictic the diploids, sexual be to appear .germanotta G. u xesv aooi apigadmlclrphylo- molecular and sampling taxonomic extensive Our G Clade— Marginata G MARGINATA AGA .intramarginalis C. .angustifolia C. .marginata C. .marginata C. nteohrhn,w see we hand, other the On . fe xmnn h yeseiesfrec of each for specimens type the examining After . .lerstenii G. Kunze, .arizonica G. and .marginata C. Gaga .monstraparva G. .marginata C. / n lorecognized also and , .siliquosa C. Kuf xLn)Ho.O h ee species seven the Of Hook. Link) ex (Kaulf. Kunth, hl l pce nteprui clade purpusii the in species all while ; HIRSUTA ru. eas ti phylogenetically is it Because group.” .Orpatdadncerphylogenies nuclear and plastid Our ). ee ftenn pce assigned species nine the of Seven hiate hirsuta Cheilanthes Fg C and 4C) (Fig. Cheilanthes and .membranacea C. Othonoloma C .hirsuta G. ru”t include to group” OMPLEX Maxon, .angustifolia C. n n eurce from resurrected one and ) ro n ro (1982) Tryon and Tryon . .hirsuta G. Cheilanthes .marginata G. .lerstenii G. Temorphological —The sioi densa Aspidotis .poeppigiana C. sascinada a as and section a as t o-ar,non- non-hairy, its ; .hirsuta C. Aspidotis and .pyramidalis C. Dvn. Maxon, (Davenp.) elne (1965) Lellinger . , .pyramidalis C. .marginata C. . (Kirkpatrick .marginata C. per to appears ehave we , sasyno- a as Gaga (Brack.) Kuhn, (see and , , ta.20;Bc ta.21) uuesuiso hscomplex this on studies Future diploids. 2010). missing putatively these al. locating on Grusz et focus should (e.g. Beck 2009; evolution al. reticulate in et of extant patterns likely critical disentangling are they are to but rare, 1). complexes or overlooked (Fig. usually species complex though nature, species cryptic this in in far Diploids so discovered individuals been diploid no have addition, In taxa. the these understand of better relationships to needed be will markers nuclear tional marginata G. “e nFg ) n fteegop ( including groups clade marginata these the of of rest the one to sister 3); as resolved Fig. in (“Me” of groups two ina hspitt eonz oeta n pce.Unfortu- species. one our than more nately, recognize to point this justifica- at tion no providing and hybridization rampant suggesting sas n ftefwpttvl ili pce nthe in species diploid putatively few 1). the (Fig. of clade it and marginata one (in scales), scales brown also dark rhizome or brown is black light lustrous have either to to contrast genus the in species oehrbtaeitrie ihother with intermixed are but together gapCp hirsuta G. of those marginata of with each G. mingle for not recovered was do allele one the Only 3). (Fig. clusters nuclear the hl h te leei oae napltm ihalleles with polytomy a in located is allele from other other with the only while forms clade one 3): well-supported Fig. in a (“Ar” alleles of types two presented it ysis, eta pmci rpod( repre- triploid locality, apomictic type an the near sent collected material including the In arrow). 4C, (Fig. surface blade abaxial the distributedon glands reddish unique with clade, marginata the in .hirsuta G. (e.g. morphology hirsuta pseudoindusial G. in extremes representing rae sasnnmof synonym a as treated aafo hsHnua pcmnt opr ihthe species. however, with distinct have, a compare do as resurrected we is to data sequence the specimen on DNA 1). Based Honduran (Fig. isotype. obtain this diploid to from sexual unable a data were be we might it Unfortunately, suggest 3315427) MO htms aecnrbtdt contributed have must as-yet-undiscovered that an and the plex of progenitor diploid sexual a between hybridization through arose that 2003) Yatskievych aooi Treatment). Taxonomic only ognize and decurrent, and pce ntemriaacaeb t non-decurrent its no Honduran has by a measurements other spore examined from measured, clade all be isotype could that from spores marginata the mature and Although the surface, pseudoindusia. blade in abaxial species the on 2, (Figs. glands distinct phylogenetically be 3). to here shown is (2004), G G G lhuhtepatdsqecsfor sequences plastid the Although aalerstenii Gaga aamarginata Gaga G MEMBRANACEA AGA LERSTENII AGA ARIZONICA AGA .spiculata C. .marginata G. lee,adteallsfrec niiuld o group not do individual each for alleles the and alleles, r setal dnia acr identical essentially are pcmn htw eune hwtoo more or two show sequenced we that specimens en h othairy, most the being gapCp notorcpoal oohltcgop;addi- groups; monophyletic reciprocally two into gapCp .hirsuta G. side eaaeett.Alsvno the of seven All entity. separate a indeed is gapCp .spiculata C. scmltl negain,wt ahname each with intergradient, completely as ifr from differs Teioyeof isotype —The Ti soeo h otdsiciespecies distinctive most the of one is —This aarcvrsm nomtv phylogenetic informative some recover data and pcmn n,ipraty hs alleles these importantly, and, specimens .lerstenii G. aacudntresolve not could data — lee htaentsse ooeanother one to sister not are that alleles .hirsuta G. n yoyieteohrto(see two other the synonymize and , aamembranacea Gaga aamembranacea Gaga .arizonica G. h otsiuae.Hr erec- we Here spiculate). most the euiu ln character. gland unique he .arizonica G. n .hirsuta G. 09( 8099 .Oursamplesof =2 .hirsuta/marginata G. .arizonica G. s l pcmn Fg 2), (Fig. specimens all oss .pyramidalis C. n .lerstenii G. aalCaldero Rafael 0 ida and Windham 90; = yMce n Smith and Mickel by G . Fg ) suggesting 3), (Fig. Fg D steonly the is 4D) (Fig. .arizonica G. hirsuta nlcigreddish lacking in .marginata G. ebaae 2 membranacea pcmn have specimens .hirsuta G. hc was which , lk species -like .arizonica G. individuals, gapCp .lerstenii G. ´ nC.066 h least the alleles, com- anal- and and )is , , Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ot iata ruswith groups that Rica Costa Y.H. NY). 86.E. 7846). eur ute td oconfirm. would to this study but further losses, require multiple by followed duplication gene 1 membranacea 37 [Volume BOTANY SYSTEMATIC 852 rohasi – otria emns n ro nCt esl eds lns etclbr m oiotlbr m iedaig modified drawings Line cm. 1 = bars horizontal mm; 1 = bars Vertical glands. reddish permission. sessile with to (2004), C in Smith arrow and and Mickel segments, from terminal to G–I in arrowheads ikl et ot eo. H.C. PH). berol., Hort. Left: Mickel. G Fig. G GERMANOTTA AGA .Mrhlg fselected of Morphology 4. .decomposita G. .pellaeopsis G. uhapeoeo a etersl fa of result the be may phenomenon a Such . ( nesnadLsosi4394 Laskowski and Anderson ( rnl 2588 Pringle (F IG )Ti sanwseisfrom species new a is 5)—This . .arizonica G. Y.I. NY). , .membranacea G. Gaga pce.A. species. .chaerophylla G. ( etyadAgels18163 Arguelles and Gentry Y B85.F. DB7815). NY; , .marginata G. norplastid our in ( ikl6254 Mickel ( rd 847 Brade .apiacea G. Y.Bakarwed nA ,adDpitt eurn suonui,grey pseudoindusia, decurrent to point D and B, A, in arrowheads Black NY). , Y.D. NY). , Y.B. NY). , mtra aet n nudtce euldpodcytotype diploid sexual of undetected an nuclear and between parent) hybrid the (maternal triploid a In is it that 2). suggesting Fig. germanotta in G. (“G” analysis ( eo 7716 Nesom .membranacea G. .marginata G. .hirsuta G. Y ooyeo h pce) G. species). the of holotype NY; , lee peri he oain “”i i.3), Fig. in (“G” locations three in appear alleles (Right: ( ikl7057 Mickel ptra aet.Temrhlg of morphology The parent). (paternal itn6837 Hinton Y enLbDtbs ceso number: accession Database Lab Fern NY; , L stp of isotype LL; , .angustifolia G. hiate spiculata Cheilanthes gapCp .membranacea G. ( ikl4782 Mickel analysis, , Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. .Aailve fsget .Rioesae etclbr=5c;hrzna as=1m.Ilsrtdb neJohnson. Anne by Illustrated mm. 1 = bars horizontal cm; 5 = bar Vertical scale. Rhizome D. segment. of view Adaxial C. rgnlyietfe as identified originally uaieprns(e aooi ramn) n n of one and Treatment), ( specimens Taxonomic paratype the (see parents putative germanotta G. 853 FERNS CHEILANTHOID OF GENUS NEW A GAGA, AL.: ET LI 2012] Fig. 5. aagermanotta Gaga smr-rls nemdaebtenistwo its between intermediate more-or-less is hiate marginata Cheilanthes ( ohese l 2618 al. et Rothfels mt Be & Smith ´ i 2081 liz UE.A ai.B bxa iwo emn.Arwedpit oacnpcosvi ending. vein conspicuous a to points Arrowhead segment. of view Abaxial B. Habit. A. DUKE). , O370)was 337501) MO , yA .Smith R. A. by ersn e pce.Atog t lsi eune are sequences plastid its Although species. new a represent n usqetyanttdas Yatskievych. and annotated subsequently and G G SP AGA 07Ti pcmn lofo ot ia may Rica, Costa from also specimen, 8097—This . .membranacea C. yMoran by Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 5 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 854 hrfr ra hsa yoy of synonym a as this treat therefore var. to ( pcmna e pce n,tu,dtriaino its of determination thus, study. and, further species awaits this status describe new taxonomic formally a to mea- as hesitant spore are specimen for we available such, were were As sporangia surement. intact we no and nuclear Unfortunately, men, obtain to pseudoindusia. unable decurrent and nteohrhand, other con- is the distribution Oaxaca). On and its Guerrero, (Jalisco, and Mexico 2), southwestern to (Fig. fined phylogeny plastid the in clade well-supported a form are isotype, the including species specimens, two distinct. geographically the and similarity, phylogenetically morphological their Despite papillate. conspicuously and thin are margins pseudoindusial in granular; seemingly slightly to smooth and feature the In thickened authors. of additional previous more margin by an overlooked The with provides 4E-F). proximally) (Fig. pseudoindusia segments (quadripinnate ultimate of divided elongate blades more leaf the primarily be that based observation species the two on the and distinguished Mickel (2004) confused. Smith easily and similar morphologically are Fg ) h stp of isotype whereas The 1). measure- diploid, (Fig. spore sexual our a species, is distinction each possible another for suggest ments specimen one on based only Although tripinnate-pinnatifid.” to tripinnate quadripinnate vs. (to dissected more blades and castaneous), vs. fro hair-tips, kinky with flexuous indicating contrast, “ clearest that the provide Mickel 189) 3). 2, (2004: (Figs. Smith species and sister are nuclear they that and indicate plastid datasets the both and similar, morphologically are foecp nsm populations. some in copy duplica- one in gene of that occurred is have groups may allele where tion two 3), these (Fig. for data explanation nuclear of the groups by two supported clearly is but purpusii G. hzm cls essetta tmyrpeetahybrid a represent may with it that suspect between brown light We and scales. pseudoindusia rhizome entire having in species that of those to identical hc pmxspeoiae,tefu pce fthe of species four ( the clade predominates, purpusii apomixis which of to specimens pled (compared entire decurrens is G. differs margin it in pseudoindusial pebbly but the pseudoindusia, that decurrent in strongly having in rd ihlgtbonmrisol n elwd.Ti clade Mexico. bicol- This to wide. if endemic cell entirely or, one is only dark margins brown uniformly light and are with blades ored, that leaf scales pentagonal rhizome include acicular clade this for characters hintoniorum G. .complanata G. G G G upsiClade— Purpusii G G APIACEA AGA PURPUSII AGA DECURRENS AGA COMPLANATA AGA .germanotta G. .harrisii C. complanata .membranacea G. .marginata G. sntrcvrdi h lsi hlgn Fg 2), (Fig. phylogeny plastid the in recovered not is sasxa ili pce Fg )weestesam- the whereas 1) (Fig. species diploid sexual a is 13i i.2 snse mn ape assigned samples among nested is 2) Fig. in 8103 pert esxa ilis(i.1.Defining 1). (Fig. diploids sexual be to appear ) /G. .purpusii G. ifr nisrioesae en somewhat being scales rhizome its in differs Temnpyyo h uaieydiploid putatively the of monophyly —The .purpusii G. .marginata G. and ) Ti pce ssmlrto similar is species —This ntepatdaayi Fg ) n we and 2), (Fig. analysis plastid the in PELLAEOPSIS ncnrs oteohrtrecae in clades three other the to contrast In / .germanotta G. .apiacea G. HARRISII .Orsoemeasurem spore Our ). hiate complanata Cheilanthes .decurrens G. tesuc ftepatdshared plastid the of source (the gapCp .purpusii G. lee r eovd plausible A resolved. are alleles — , pert eaoitctriploids. apomictic be to appear — .apiacea G. aacomplanata Gaga dae akr(atropurpureous darker axes nd lhuhitrige with intermingled although , .harrisii G. aaapiacea Gaga eune rmti speci- this from sequences .complanata G. Fg ) ti itntfrom distinct is it 2), (Fig. hc a iia entire similar has which , .apiacea G. ihsbeun loss subsequent with , , , a etetraploid be may .pellaeopsis G. .apiacea G. that and .pellaeopsis G. nssgetthat suggest ents hsmri is margin this , aapellaeopsis Gaga var. and .complanata G. . .marginata G. .pellaeopsis G. .harrisii G. interrupta edto tend and , the , .decomposita G. Morphologically, 3). Fig. in “D” and chaerophylla G. ( 3). the groups Fig. In two or 2). one (Fig. either the phylogeny and clade, all plastid well-supported 3), the (Fig. phylogeny in nuclear reflected observa- is this and tion complex,” the of bound- members sharp no the be between to aries seem “there that noted (2004) Smith and Mickel prevalent. are hybridization and polyploidy, apomixis, ti uefcal oesmlrto cm). 3.2–6.5 similar vs. more (15 superficially pinnae basal is well longer as It much 4-pinnate proximally, are having 3-pinnate) that as or blades 2-pinnate-pinnatifid leaf lanceolate) (vs. (vs. ovate having separated. of Although origin 3). “Mo” between parent; Fig. paternal hybrid in (putative species a diploid unknown be an to cuneata appears G. Treatment), Taxonomic in angustifolia G. (as segments as tified tripinnate terminal with angustifolia linear-lanceolate G. G-I) and 4 in conform (Fig. (as leaves progenitors tripinnatifid to two its between ihisifre eoi osiuin(w eoe from genomes (two constitution angustifolia genomic to G. inferred closer its much with is specimen this the to similar both on the visible in as are Here, non- surfaces. that blade spores, veins tan) and or pseudoindusia, brown decurrent than (rather black have members i.3.Ti ugssthat suggests the This from 3). one Fig. and 3) Fig. in is fpplae.Ti pce a nybe eotdfrom reported been only has species Leo Nuevo This spe- other papillate). all if in cies, (irregular protrusions papillate margins regular pseudoindusial segments, The its have blades. elongate with leaf deltate scales, genus, strongly rhizome and the acicular in of species combination distinctive and recognizable lds ti ekyspotda itrt h angustifolia the to sister as supported leaf and weakly in rachises, is genus petioles, the It on in blades. unique hairs is glandular it dense and having clade, this to assigned rently collection, sterile one unknown. only is from level pinnules. ploidy known its acroscopic and is the species than new longer This slightly only basiscopic basal pinnules and blades leaf deltate-pentagonal) (vs. ovate upre ld ntencerpyoeyta sdsic from distinct is pellaeopsis that G. phylogeny nuclear the in clade supported purpusii G. rmZrgz,NeoLeo Nuevo Zaragoza, from apiacea Leo (Nuevo Mexico northeastern to eadtonwrcrsfo aalps oho hc were as which misidentified of previously both Tamaulipas, from records new two add we aamonstraparva Gaga nutflaClade— Angustifolia alusiClade— Kaulfussii G G HINTONIORUM AGA nutfla1 angustifolia .angustifolia G. a ecie nyrcnl,bsdo w collections two on based recently, only described was aadecomposita Gaga .angustifolia G. .monstraparva G. .decomposita G. ´ pttv aenlprn;“o nFg.2 )and 3) 2, Figs. in “Mo” parent; maternal (putative ,Mxc,o ysmsubstrates. gypsum on Mexico, n, ntepatdpyoey(i.2,frsawell- a forms 2), (Fig. phylogeny plastid the in aamonstraparva Gaga Fg ) h itiuinof distribution The 3). (Fig. i.4G ryarwed) n pcmniden- specimen One arrowheads). grey G, 4 Fig. , n n from one and oeec from each (one u ifr rmbt hs pce nhaving in species these both from differs but pttv aenlprn;“”i is ,3) 2, Figs. in “D” parent; maternal (putative ru.Itrsigy h opooyof morphology the Interestingly, group. ( nutfla2 angustifolia — e pce ecie ee(see here described species new a , .cuneata G. aakaulfussii Gaga 64)hsthree has (6541) aahintoniorum Gaga u iha diinlgnm from genome additional an with but hscaei nqei htall that in unique is clade This h w pce r ahreasily rather are species two the , ssont eahbi between hybrid a be to shown is ´ nutfla1 angustifolia .pellaeopsis G. .chaerophylla G. Mce n mt 04.Here 2004). Smith and (Mickel n .angustifolia G. .chaerophylla G. .chaerophylla G. aamarginata/hirsuta Gaga nutfla1 angustifolia .angustifolia G. .chaerophylla G. .decomposita G. spttv aenlparent; paternal putative as , ifr from differs a lal novdi the in involved clearly was ´ .angustifolia G. n Tamaulipas). and n steol pce cur- species only the is gapCp .apiacea G. .chaerophylla G. Apni 1). (Appendix and sprastemost the perhaps is ). i.4) e with yet 4I), Fig. , 51i hybrid a is 6541 and lee,tofrom two alleles, ru “n in (“An” group nutfla2 angustifolia lee alinto fall alleles sintermediate is .cuneata G. lee oma form alleles nutfla2 angustifolia congruent , srestricted is complex, Gaga and in in Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. otwsenUie tts(rzn n ea)t south- to Texas) Bolivia. and ern (Arizona States United southwestern ubrpresumably number (as sal lbosbae ecp npedidsai several in surfaces in blade pseudoindusia and and sporangium, species on per (except large) blades 16 glabrous or or small usually 32 small (vs. 64 spores large of 32 production the pseudoindusia, differ- inframarginal surfaces; from blade adaxial ing striate and lustrous inently rae rsiuae prni 2soe r64-spored. numbers: or Chromosome black. 32-spored or fim- n brown Sporangia in papillate, tan, globose; rachises) pebbly, spiculate. Spores entire, and or margins hairy; costae briate, the to along species, glabrous extending some Pseudoindusia (i.e. tips. decurrent vein at sporangia the clustered endings pseudoindusia, strongly vein by inframarginal the protected differentiated Sori elongate, hydathodes. to prominent linear forming often for oblong, costules (except ovate, and glabrous Segments basiscopic usually costae basal adaxially; the grooved the exaggerated, pentagonal, usually to pinnules to 2- Blades ovate-deltate scales. occa- hair-like 4-pinnate, adaxially, or hairs grooved black, scattered usually with Rachises sionally to base. at castaneous in scaly in ally Petioles hairy brown (glandular margins. (pale glabrous mostly brown lustrous and often stripe and acicular, black membranacea to G. to lanceolate scales brown Rhizome dark scaly. horizontal, or 02 IE L:GG,ANWGNSO HIATODFRS855 taxonomy. provide diploids, and stable boundaries sexual species a delimit of fully to morphology unable spe- are accepted the we the of knowing half Without than less cies. in found sex- were putative diploids morphological Furthermore, ual many included. and not that complete were noted from intermediates be far clarify is should and sampling it our events However, hybridization issues. past taxonomic resolve several were we partly data, to spore with able data molecular nuclear the and plastid in species vague and hirsuta particularly FERNS genus CHEILANTHOID the are OF GENUS of boundaries NEW taxonomy A the GAGA, have AL.: complicated apomixis ET greatly LI and polyploidization, Hybridization, unfold. 1). (Fig. triploids apomictic be clades. to purpusii inferred of are non- or study specimens marginata have the two clades in The both seen of character not a members scales, combination rhizome and concolored and 3) indusia decurrent 2, (Figs. clade 2012] agn fmtr evs(s moha 40 at bullate smooth (vs. minutely leaves mature apices, of segment margins mucronate) (vs. attenuate Gaga efforts. these to (or critical be loci will nuclear data) microsatellite additional incorporating and Identifying diploids revision. taxonomic sexual careful and study systematic =2 Distribution— lnstretil hzmssotadcmat ascending compact, and short Rhizomes terrestrial. Plants nwr otemseisof mysteries the to Answers otsmlrto similar Most Cheilanthes hiate marginata Cheilanthes marginata n re,FyWiL,&Wnhm gen.nov.—TYPE: Windham, & Li, Fay-Wei Pryer, 0(pmci rpod in triploid) (apomictic 90 = ope n nteagsioi ld.B combining By clade. angustifolia the in and complex Cheilanthes aamarginata Gaga Knh a-e i&Wnhm Basionym: Windham. & Li Fay-Wei (Kunth) nWnhmadYtkeyh20) h base the 2003); Yatskievych and Windham in ,cnooe,o iooe ihdr central dark with bicolored or concolored, ), h itiuinof distribution The aooi treatment Taxonomic Aspidotis Gaga x =30. .s nissrnl differentiated, strongly its in s. s. .kaulfussii G. sapiecniaefradditional for candidate prime a is Kunth u ifrn nisruddto rounded its in differing but aakaulfussii Gaga stems ieydistributed widely most the is Gaga .arizonica G. .kaulfussii G. nlddi u spore our in included Gaga aejs eu to begun just have

). + agsfo the from ranges ,adls prom- less and ), aamarginata/ Gaga and .kaulfussii G. ,occasion- ), .cuneata G. Gaga ). fmcocpcadmcocpcfaue sediagnosis (see features from separate to impossible microscopic nearly is it and above), macroscopic genus of sister its from guished of species CAGG]. fern related has type cheilanthoid the closely other and any the GAGG, in site [e.g. this sampled at seen not pattern name The indi- work. of our diversity the fund to who relevant find- viduals and our accessible make to more and ings interests, public confluence and the acknowledge science of to between research, us allows patterns basic honor supports naming cryptic this funding underlying public Because biology diversity. under- intricate to a the struggle with the stand her Gaga characterizes differ- provide that hereby broad Lady namesake we celebrate scientific species, to Because own humanity its society. within for ences need individ- today’s the and to in equality speaks of expression defense fervent ual and articulate for Gaga, her Lady singer-songwriter-performer pop American eu.A uloiepstos5861i the in 598–601 positions all alignment, nucleotide the At characterizes that genus. synapomorphies molecular the of one iigeatyweet rwtebudre.Our boundaries. the nested of draw series a to identify analyses deter- where still molecular are unpublished we exactly and genera prep.), in fern mining al. et cheilanthoid of (Windham progress recircumscription other in The most nested. and are it which within found be can species endemic. are 19 these the of six of species and 17 of Mexico, in genus; center the the is for Colombia, Mexico diversity Honduras, Bolivia. Rica, and Mexico, Costa Peru, Nicaragua, from Ecuador, Salvador, recorded El Guatemala, been having species, pmrh of apomorphy opooia Affinities— Morphological Etymology— Fig. .Pr fthe of Part 6. Gaga Gaga h genus The .Only matK pce ae“AA Fg ) sequence a 6), (Fig. “GAGA” have species Gaga lgmn hwn eiigmlclrsyn- molecular defining a showing alignment pce ae“AA tpstos598–601. positions at “GAGA” have species Gaga Although Aspidotis snmdi oo fthe of honor in named is Cheilanthes sioi densa Aspidotis Cheilanthes Gaga sn combination a using Gaga sesl distin- easily is ( Cheilanthes .micropteris C. loechoes also matK sawork a is shows gene .l., s. ) Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. wle rtbrne.Terioe of rhizomes The protuberences. swollen without rhizome short-creeping or compact a having in these of .bolborrhiza C. are h ld ufcso oeo h fia pce are species African the of some of glabrous. though surfaces nearly leaves, projections scaly blade and/or hairy glassy the have the generally of conical, clade species s. contrast, s. By distinctive spiculae. a as of to or produce referred clade) blades marginata glands on leaf the the in pubescent sessile 3) notably are (most and species margins, species few and/or several surfaces of their 2) trichomes; pseudoindusia gland-tipped of the multicellular, blades with leaf in covered the glabrous are 1) largely exceptions: are following These the blades. leaf the of lineages distinguishing cell prep.). the character in Cheilanthes al. in morphological et mitosis third (Windham premeiotic cells A mother a spore of the loss generating sporangium sporan- the per per spores to large spores due 16 not produce small apomicts (but 32 and gium Australian produce species and reproducing American South of species African) the stark apomicts in to stands whereas This sporangium. sporangium contrast per per spores large spores 32 small produce 64 of ance eulyrpouigseisof species reproducing sexually another of discovery the of the in species to all led distinguishes that has feature spo- size per spore number and spore rangium on focus research Our inframarginal. in only and Aspidotis character), this by in defined resided previously (many Cheilanthes .W ie l nul data). unpubl. these al. et Li of W. to F. none related closely that are indicate species results molecular Unpublished ( sessile either on present are glands When surfaces. abaxial the rufopunctata most C. of the leaves to contrast glabrous in Furthermore, bases. root swollen prominent .glauca C. 5 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 856 tdadceryifaagnl(eesdfo h efedge) leaf the from (recessed differenti- in inframarginal strongly clearly the is and is which ated these pseudoindusium, among the Foremost of absolute. nature is which of none ters, from the of One coherence. of circumscriptions morphological defensible most of degree some show species (1982), squamosa wouldC. genus including the species American of Australian concept all revised ana- our encompass this 2B), species African Fig. that their of suggest in clade lyses Clade 1 the (Clade to (2011; authors addition these In al. by identified 2B). et Fig. their Eiserhardt in of C analyses phylogenetic the lds l fwihme h iiu eurmnsfra for requirements minimum the of meet concept revised which of all clades, Within The Gaga .rufopunctata C. Gaga Cheilanthes .obducta C. Cheilanthes n t itrgenus sister its and .micropteris C. ) r ogcepn n hs of those and long-creeping are Cheilanthes schimperi ae nacmiaino opooia charac- morphological of combination a on based .s ld aeapol eie pseudoindusium defined poorly a have clade s. s. .s sdfndaoerltst h pubescence the to relates above defined as s. s. ok rv pce ru fTynadTryon and Tryon of group species Grev. & Hook. ikl&Beitel. & Mickel .arizonica G. aerdih tle,gadlrtihmson trichomes glandular stalked, reddish, have et xKh,and Kuhn, ex Mett. .s ld.Lk otlpoprnit ferns, leptosporangiate most Like clade. s. s. .s ld ulndaoei distinguished is above outlined clade s. s. Cheilanthes Cheilanthes ostepedidsu pert be to appear pseudoindusium the does Rosenst., r esnbywl upre,and supported, well reasonably are , .l,teseisms iia to similar most species the l., s. rnta l eds ( reddish all at not or ) Gaga Cheilanthes .micropteris C. Aspidotis Gaga Gaga .s ntelte ld,sexually clade, latter the In s. s. Notholaena nta hyicuetetype the include they that in .glauca C. pce,both species, Gaga sdsigihbefo each from distinguishable is Wnhme l nprep.; in al. et (Windham .fractifera C. Cheilanthes eryalseiso the of species all Nearly . lsagopo South of group a plus rdc preponder- a produce Gaga Cheilanthes tetp pce) the species), type (the .rufopunctata C. hc fe was often which , .bolborrhiza C. Cv)Mt.and Mett. (Cav.) rmmn taxa many from .M Tryon. M. R. .glauca C. mre from emerges Gaga .kaulfussii G. .kaulfussii G. Cheilanthes Gaga Gaga (formerly hyare they , ,with from Gaga have and and ). .Gg arizonica Gaga 3. .Gg angustifolia Gaga 1. .Gg complanata Gaga 5. chaerophylla Gaga 4. .Gg apiacea Gaga 2. .Gg cuneata Gaga 6. .Gg decurrens Gaga 8. loou angustifolius Allosorus hiate venusta Cheilanthes hiate complanata Cheilanthes .Gg decomposita Gaga 7. e Combinations— New e okBt ad 8 8.20.TP:MEXICO. 2004.—TYPE: 189. 88: Gard. Guerrero, Bot. York New o.nd—YE EIO aaa a er de Pedro San Oaxaca: MEXICO. Nolasco, nud.—TYPE: nom. :1.15.TP:MXC.Oxc:Juquila, Oaxaca: MEXICO. 6362 1857.—TYPE: 11. 9: ei.Ben 1 rpit6) 89 TP:MEXICO. —TYPE: 1849. Yavesia, 67). Oaxaca: (reprint 219 Bregn. Mexic. n. s. Bonpland ad 8 9.20.TP:GAEAA Volca GUATEMALA. 2004.—TYPE: 191. 88: Gard. nov., nov., 7716 Nesom 1981). 71. 2: Chiapas Fl. in Sm. R. A. Michoaca MEXICO. 1867.—TYPE: 150. Farrnkra angustifolia Kunze, 1836. 152. (Kunth) Pterid. Tent. Suppl. p :2.1815. 21. 1: Sp. nov., comb. Agua, aeti ok,S.Fl :11 1852. 151. 2: Fil. Sp. decomposita Hook., Galeotti) MO!). MEXU, de Motozintla Mendoza, Chiapas: MEXICO. 1980.—TYPE: 19. 70: nov., comb. Oaxaca: MEXICO. 1850.—TYPE: 307. Juquila, 243, 23: Linnaea chaerophylla Me Cheilanthes Galeotti, & nov., comb. Windham, m enJ :16 1918. 116. 8: J. Fern Am. 8 8.20.TP:MXC.NeoLeo Nuevo MEXICO. 2004.—TYPE: 182. 88: 8 l 0 .2 1842. 2. f. 10, pl. Me 48, Nouv. Galeotti, & ida,cm.nov., comb. Windham, PH). isotype: B; (holotype: plants cul- tivated from Described 1910.—TYPE: 5. 51: Brandenburg Prov. angustifolia cuneata cuneata cuneata Cheilanthes 99—YE .S .Aioa ucuaMts, Huachuca 1327 Arizona: A. S. U. 1979.—TYPE: arizonica Cheilanthes arizonica nov., (BR). hltp:U! stp:NY). isotype: US!; (holotype: hiate apiacea Cheilanthes hiate pyramidalis Cheilanthes hiate decurrens Cheilanthes ao a 3709 Hay & Maxon Ln)J m,J o.(ok)4 159. 4: (Hook.) Bot. J. Sm., J. (Link) aeti6367 Galeotti Ln)J m,Hs.Fl 8.1875. 286. Fil. Hist. Sm., J. (Link) si .149 M. Espin Mxn ru,IdxN mr en5.1938. 51. Fern Amer. N. Index Broun, (Maxon) aeti6560 Galeotti ut var. Kunth hltp:N! stps A! TEX). CAS!, isotypes: NY!; (holotype: Knh ae nHoe ae,Sn Fil. Syn. Baker, & Hooker in Baker (Kunth) relv 41747 Breedlove M atn aeti Fe Galeotti) & Martens (M. hiate complanata Cheilanthes Ln)FyWiL ida,cm.nov., comb. Windham, & Li Fay-Wei (Link) hiate angustifolia Cheilanthes Mce)FyWiL ida,comb. Windham, & Li Fay-Wei (Mickel) BWld 01,P-Humb.). 20116, (B-Willd. Fe Mxn a-e i&Wnhm comb. Windham, & Li Fay-Wei (Maxon) Mce)FyWiL ida,comb. Windham, & Li Fay-Wei (Mickel) ´ ´ loou angustifolius Allosorus var. e .Fu.Mxqe4,p.1.1842, 11. pl. 47, Mexique Foug. m. A .S. a-e i&Windham, & Li Fay-Wei Sm.) R. (A. ibans n. s. Liebmann M atn aeti a-e i& Li Fay-Wei Galeotti) & Martens (M. ..S.var. Sm. A.R. M atn aeti a-e i& Li Fay-Wei Galeotti) & Martens (M. Knh .Pelvar. Presl C. (Kunth) ik ot eo.2 0 1833. 40. 2: Berol. Hort. Link, Knh a-e i&Windham, & Li Fay-Wei (Kunth) Mxn ikl htlga4:433. 41: Phytologia Mickel, (Maxon) hltp:N! stps NB FCME). ENCB, isotypes: NY!; (holotype: hltp:B;ioye P). isotype: BR; (holotype: ele decompositus Pellaea ´ (BR). aurata ikl e.NwYr o.Gard. Bot. York New Mem. Mickel, cuneata .Aa.Ry c.Buels15: Bruxelles Sci. Roy. Acad. m. loou chaerophyllus Allosorus loou decompositus Allosorus M atn aeti Kunze, Galeotti) & Martens (M. hiate pyramidalis Cheilanthes (NY!). ikl e.NwYr Bot. York New Mem. Mickel, ¨ tr2 1 1848. 11. 2: uter Fe Fe hltp:D! isotypes: DS!; (holotype: ´ ´ rue eh o.Vereins Bot. Verh. Brause, subsp. e ,Me e, lcoye ,coe by chosen C, (lectotype: interrupta ncimangustifolium Onychium .R m,A.Fr J. Fern Am. Sm., R. A. ´ .Fu.9 2 1857, 12. 9: Foug. m. ut,Nv Gen. Nov. Kunth, Knh .Presl, C. (Kunth) arizonica ´ ´ n, ,Me e, M atn & Martens (M. minus ikl Mem. Mickel, ´ :Zaragoza, n: ublt& Humboldt 1841. .Martens M. .Martens M. Cheilanthes Cheilanthes ´ Cassebeera .Foug. m. Goodding Fe Maxon, Liebm., Galeotti ´ Pellaea Pellaea ´ var. e nde Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 2 aakaulfussii Gaga 12. hintoniorum Gaga 11. angustifolia Pellaea hiate glandulifera Cheilanthes 4 aamarginata Gaga 14. 3 aalerstenii Gaga 13. hiate selinoides Cheilanthes membranacea Gaga 15. loou ciliatus Allosorus .Gg harrisii Gaga 9. hiate spiculata Cheilanthes pyramidalis Cheilanthes 6 aapellaeopsis Gaga 16. 0 aahirsuta Gaga 10. hiate rufescens Cheilanthes 02 IE L:GG,ANWGNSO HIATODFRS857 FERNS CHEILANTHOID OF GENUS NEW A GAGA, AL.: ET LI 2012] hiate crenulata Cheilanthes haa:Ms eCoapilla, de Mesa Chiapas: trdg.SrMe Sur Pteridogr. 6837 al. et Hinton 89—YE EIO Oaxaca, MEXICO. 1849.—TYPE: 9 85—YE MEXICO. 1825.—TYPE: 59. 1.20.TP:MXC.Me MEXICO. 2004.—TYPE: 210. Agostı San MEXICO. Guatimalpan, MEXICO. P); Mexico, of Valley MEXICO. TYPE: 2 1981). 72. C;isolectotype:K,chosenbyA.R.Sm.,Fl.Chiapas2: 92—YE EIO aic:Nvd eColima, de Nevado 12869 Jalisco: McVaugh MEXICO. 1992.—TYPE: ecie rmcliae plant cultivated from Described ecie rmcliae plant cultivated from Described nov., UC). NY, MEXU, NuevoLeo MEXICO. 1991.—TYPE: 1, f. 551, 14: Sida Nesom, & lns(,BM). (B, cultivated plants from Described 1839.—TYPE: 145. 13: Linnaea 1833. 43. 2: Berol. Hort. 1818, 1815, nov., ida,cm.nov., comb. Windham, stp:BWld 20126). Quito, B-Willd. n. isotype: s. ECUADOR. Bonpland 1865.—TYPE: ao,A.Fr .8 1.11.TP:MEXICO. 1918.—TYPE: 119. 8: Felipe, San J. de Sierra Fern Oaxaca: Am. 1896. 6. Maxon, 5, f. 18, t. 262, nov., comb. 4 1 92—YE JAMAICA. 1922.—TYPE: 51. 24: nov., ae lns(! ebru acd:B2 0036984, 20 B barcode: 1832’). Berol. herbarium ‘Hort. as culti- (B!; labeled from plants Described vated here): (designated LECTOTYPE hirsuta Cheilanthes aaa oad Vega, UC!). isotype: de MEXICO. 1988.—TYPE: Sola 117. 46: Oaxaca: Gard. Bot. York New nov., comb. ob nov., comb. ´ :Galeana, n: hiate harrisii Cheilanthes hiate marginata Cheilanthes hiate viscosa Cheilanthes ele marginata Pellaea .Pelvar. Presl C. hiate pellaeopsis Cheilanthes ele membranacea Pellaea hiate lerstenii Cheilanthes Ln)FyWiL ida,cm.nov., comb. Windham, & Li Fay-Wei (Link) Mxn a-e i&Wnhm comb. Windham, & Li Fay-Wei (Maxon) Mce etl a-e i&Windham, & Li Fay-Wei Beitel) & (Mickel itne l 18765 al. et Hinton ikli cag,F.Nv-a.1:243. 17: Nov.-Gal. Fl. McVaugh, in Mickel hltp:MC;ioye US). isotype: MICH; (holotype: Knh ae var. Baker (Kunth) Mickel,Mem.NewYorkBot.Gard.88: Kne a-e i&Wnhm comb. Windham, & Li Fay-Wei (Kunze) Knh a-e i&Wnhm comb. Windham, & Li Fay-Wei (Kunth) hltp:P ihrslto photo!; resolution high P, (holotype: ik ot eo.2 9 1833.—TYPE: 39. 2: Berol. Hort. Link, ik ot eo.2 1 1833.—TYPE: 41. 2: Berol. Hort. Link, hltp:N! stps ,L! MO!). LL!, F, isotypes: NY!; (holotype: Fe ib. ei.Brn 5 rpit106). (reprint 258 Bergn. Mexic. Liebm., ´ ´ Mce)FyWiL Windham, & Li Fay-Wei (Mickel) Mneh eo)FyWiL & Li Fay-Wei Nesom) & (Mendenh. ,Me e, ´ io10 99—YE MEXICO. 1909.—TYPE: 130. xico ik ot eo.2 0 1833.— 40. 2: Berol. Hort. Link, Dvn. a-e i&Windham, & Li Fay-Wei (Davenp.) n, hiate membranacea Cheilanthes cafe 305 Schaffner hiate hintoniorum Cheilanthes ´ ao,Cnr .S al Herb. Natl. S. U. Contr. Maxon, iknm le. o Carmich non illeg., nom. Link Knh ae,Sn i.151. Fil. Syn. Baker, (Kunth) .Fu.7 8 .2,f .1857.— 3. f. 25, t. 38, 7: Foug. m. minor ikl6238 Mickel ut,Nv e.S.1 22. 1: Sp. Gen. Nov. Kunth, oioa1059 Rovirosa ak .n. s. Haeke hiate kaulfussii Cheilanthes cafe 304 Schaffner rnl 5963 Pringle hltp:B stp:PH!). isotype: B; (holotype: s hltp:B stp:PH!). isotype: B; (holotype: s .Pel ei.Hek 1: Haenk. Reliq. Presl, C. hltp:TX isotypes: TEX; (holotype: ibans n. s. Liebmann ars12902 Harris ikl&Bie,Mem. Beitel, & Mickel aep,Bt a.21: Gaz. Bot. Davenp., ikli cag,Fl. McVaugh, in Mickel ´ io Temascaltepec, xico: cafe 88 Schaffner ioytp:K). (isosyntype: elongata P rPRC). or (PR hltp:NY; (holotype: P,fa.US). frag. (PH, (GH). ublt& Humboldt snye P!); (syntype: (US). (lectotype: Mendenh. Rovirosa, (Davenp.) (syntype: Kunze, lc,cnooe rwt ihe ae eilst 0cm 30 to lustrous, Petioles base. lanceolate, lighter wide; a mm with long 1 or to concolored long, black, mm 7–10 scales coriaceous somewhat venation. and obscure with blades leaf scales, thin-textured) (vs. rhizome castaneous) (vs. to mar- similar brown gins; obvious lacking scales rhizome black concolored, 9593 8.64,334meeain 3Jn 2008, June 23 Chirripo Muerte, al Gerardo 2618 elevation, la San al. de et m de Rothfels Cerro J. Tapanti-Macizo 3,334 C. Sendero National W83.76144, of Parque N9.57983, beginning of Estaqueros, edge Cerro at de pie Sakira, Panamericana, Sabila-Cerro Zacatales-Cerro of side brown. globose, 5. Figure Spores unknown. one-celled number Chromosome 32-spored. irregular with Sporangia margin protrusions. costules; strongly along surface; segment decurrent abaxial the half Pseudoindusia than obscure. long, more veins covering linear; mm to oblong 3–8 wide, mm segments 1 longer ca. Ultimate slightly pinnules. non-conform, pinnules acroscopic lobed, basiscopic than with basal to often segment; deltate cm pinnules terminal 6 coriaceous; long, and cm pinnae 9 somewhat ca. wide; pinnae wide, largest 4-pinnate; cm to to blades 3- 11 pentagonal, Leaf long, glabrous. groove, cm of 15 sides with both adaxially, on grooved ridges Rachises pale base. at scales linear sparse t2,K 78,c.1k Wo Asuncio of NW km 1 Hwy, ca. American 87–88, (Pan Km General 2), del Rt Isidro San to Cartago from 129 Mehltreter K. moe ot n npr rvr nodrt aethe make to order to place. in braver is bravery kinder, a mission inspire are world Its and differences celebrated. youth where is empower organization society, individuality and accepting charitable embraced more non-profit a to fostering new dedicated is foundation a The (bornthiswayfoundation.org). & Founda- (BTWF), Way This D. Born tion Society, the John & launched family Internet Germanotta for The the Center Berkman with Endow- Harvard’s and California partnership On ment, The Foundation, Germanotta. in MacArthur Joe T. 2012, Catherine and Cynthia 29, parents, February Gaga’s Lady of Chirripo Cerro and Muerte la de Cerro in ,0 lvto,o te odbn.2 a 1986, Jan 29 bank. road Be T. steep & 2018 on Smith elevation, R. A. m 3,200 tion, 7 aapurpusii Gaga 17. .Gg germanotta Gaga 1. eils( mi imtr,adlre t 10 (to larger and diameter), in mm 2 (> petioles lnstretil hzmssotadcmat Rhizome compact. and short Rhizomes terrestrial. Plants Paratypes— Etymology— Distribution— e Species— New otsmlrto similar Most ob nov., comb. 92—YE EIO a usPotosı Luis Rafael, San MEXICO. 1982.—TYPE: UE stps R N,M,U,U,K NY). N9.57983, K, US, UC, 2012, MO, Cerro INB, June CR, isotypes: Muerte, DUKE; 18 elevation, m la W-facing 3334 vegetation, W83.76144, Sakira, de Cerro paramo to Cerro slope, trailhead the 2, near of Zacatales, NE-side Hwy Cartago, de Panamerican Provincia RICA. COSTA TYPE: + Jalisco: MEXICO. 1992.—TYPE: 241. Zapotitla 17: Nov.-Gal. . mi imtr atnos lbosecp for except glabrous castaneous, diameter, in mm 2.5 ups4881 Purpus ´ n, OT IA rv a Jose San Prov. RICA. COSTA aamembranacea Gaga h pte graot”i ae nhonor in named is “germanotta” epithet The hiate purpusiiCheilanthes li tal. et Iltis h pce skonol rmpa from only known is species The aamarginata Gaga C 600.Po.SnJose San Prov. 160010). (CR T evs a-e i&Windham, & Li Fay-Wei Reeves) (T. a-e i&Wnhm p nov.— sp. Windham, & Li Fay-Wei ´ hltp:G;ioye:F C,MO!). UC!, F, isotypes: GH; (holotype: DK) ua ehlag:Camino hallazgo: de Lugar (DUKE). liz 94 WS rg NY!). frag. (WIS, 29543 ´ C 284 O3337501). MO 122854, (CR ,5 lvto,2 pi 1991, April 26 elevation, m 3,250 , u ifrn nhvn thicker having in differing but u ifrn nhvn black having in differing but a-e i1520 Li Fay-Wei .Ree,Rooa8:293. 84: Rhodora Reeves, T. ´ ot Rica. Costa , ´ ,pa n, ´ ee Zeledon, Perez , ´ ´ :MinasdeSan

Crao road /Cartago, + ´ 1mm),mostly aovegeta- ramo (holotype: ´ aoof ramo Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. that unfurling. to inrolled prior tightly leaf a fern to young resemblance striking a claw” bears “monster which outstretched offi- hand, the The is monsters.” greeting “little monster little her cial fans, loyal and fervent Gaga’s yrdzto between hybridization germanotta inlsa ot15tmslne hnarsoi pinnules. acroscopic 18 than longer to basiscopic segments times basal Ultimate 1.5 segment; most terminal at conform, pinnules a with pinnules obnMrnadJfesnPaofrcaiyn h aooi ttsof status to taxonomic the respectively; clarifying UC, to for and permission Prado Othonoloma NY Jefferson for at and specimens Smith Moran type Alan Robbin from are and samples We Moran DNA Mickel. Robbin take John to by laid grateful groundwork especially taxonomic careful the without otnospoial;mri oeo esetr and entire less or unknown. number more Chromosome and unknown. Spores margin smooth. adaxially the proximally; of dis- both occasionally continuous one-third and non-decurrent visible to surface; segment half abaxial veins covering Pseudoindusia secondary abaxially. and mary n iial iegn from divergent minimally 3), one of (Fig. samples dataset three the nuclear to sister of plastid as accession supported the only the In 2), clade. (Fig. angustifolia dataset the from diploid unknown 5 YTMTCBTN Vlm 37 [Volume BOTANY SYSTEMATIC 858 o)ada neetdsxa ili yoyeof cytotype diploid sexual undetected an and tor) nfiitdwt n ftewl-upre rusi the in groups well-supported the clade. of angustifolia any with unaffiliated h ag,lsru,bakrioesae of and both scales) brown scales from concolored, it rhizome distinguish black immediately lustrous, large, The a 6c og 6c ie oehtcraeu,ovate, coriaceous, somewhat wide; blades 15 cm ca. Leaf pinnae 16 glabrous. largest proximally, long, groove, 4-pinnate of cm sides 36 both with ca. adaxially, on diame- grooved ridges in mm Rachises pale 2.7 glabrous. and long castaneous, cm ter, concolored, 35 lanceolate, ca. wide, Petioles mm black. 0.7 lustrous ca. long, mm 3–4 scales blades. leaf on scales of lack complete a lin- and (vs. ear), lanceolate narrowly segments the continuous, inantly to similar only pinnules; acroscopic pinnules angustifolia the basiscopic than basal longer and slightly blades leaf pentagonal) decomposita G. ine(5v.3265c) iia to similar cm); basal 3.2–6.5 longer vs. 2-pinnate- much (15 (vs. and proximally 4-pinnate pinnae 3-pinnate) are or that pinnatifid blades leaf lanceolate) monstraparva Gaga 2. narrow diagnosis. and pro- species the stripe are in features vided central distinguishing dark Additional a margins). brown with scales bicolored ally Relationships— Etymology— Distribution— Acknowledgments. Relationships— lnstretil hzmssotadcmat Rhizome compact. and short Rhizomes terrestrial. Plants iia to Similar 9.37,225meeain 9Ags 2009, August Ocuila 1229 09 elevation, m de 2,295 W99.33072, Me Municipio Ocuila of on west km Mexico, 3.9 approximately MEXICO. TYPE: .monstraparva G. oGria ohnfrtkn ihrslto htso the of photos resolution high taking for Rouhan Germinal to ; hltp:DUKE). (holotype: u ifrn nhvn h suonui predom- pseudoindusia the having in differing but sa pmci rpodta rgntdthrough originated that triploid apomictic an is aacuneata Gaga ´ eAtaaCenvc od N18.98474, road, Arteaga/Cuernavaca de n h pte “ epithet The u ifrn nhvn vt v.deltate- (vs. ovate having in differing but hsseisi nykonfo h holotype. the from known only is species This u hlgntcrcntutossuggest reconstructions phylogenetic Our pr n N aasgetthat suggest data DNA and Spore sahbi involving hybrid a is hssuycudnthv encompleted been have not could study This .membranacea G. a-e i&Wnhm p nov.— sp. Windham, & Li Fay-Wei .monstraparva G. + u ifrn nhvn vt (vs. ovate having in differing but m arwylnelt;pri- lanceolate; narrowly mm, 3 .marginata G. monstraparva” .cuneata G. .monstraparva G. temtra progeni- maternal (the ´ .membranacea G. ioMrlsborder xico/Morelos .chaerophylla G. hw w alleles, two shows

+ usu- smaller, (with .cuneata G. .cuneata G. 8cm;pinnaeand n h other the and oosLady honors .germanotta G. .marginata G. .B Beck B. J. swell- is nthe In . n an and (with Gaga Gaga and ´ n, . aa oAn ono o h lutainof illustration the the for interpreting Johnson on Anne discussions for to Schuettpelz data; Eric to Rica; Costa ek .B,M .Wnhm .Ytkeyh n .M re.21.A 2010. Pryer. M. K. and Yatskievych, G. Windham, D. M. B., J. Beck, by part Jose of Ded- in memory MDW. the and funded to KMP icated to was DEB-0717398 grant research Foundation Science This National invaluable. were course encouragement biology Franc systematic by a taught for in Duke undertaken at Grusz project Amanda term study a and This from assistance. Sigel, laboratory blossomed and Erin manuscript the Huiet, on Layne feedback valuable wish Rothfels, (Research also Carl Bates We thank Karl obvious. the to to stating and for Barrie Latin, Univeristy) Stud- on for Fred Duke (Classical guidance Communications, PH to Atkins for and grateful Jed University) to are B, Duke advice, ies, We CAS, nomenclatural study. F, for this TEX/LL, Museum) (Field to MO, critical NY, specimens DUKE, loaning of curators and for staff primers sequencing providing for Huiet iehrt .L,J .Rhe,S .Rsel .C eiyr,and Yesilyurt, C. J. Russell, J. S. Rohwer, G. J. L., W. Eiserhardt, hitnuz .J . .C hn,adH cnie.21.Alinear A 2011. 1906. C. Schneider. Christensen, H. and Zhang, C. X. M., J. M. Christenhusz, MO Carl visit and Beck to their James to FWL access to providing supporting DEB-0717430); for for Rothfels Foundation Yatskievych Science George (National to treatment; marginata Gaga oaa .adK .Cadl.19.MDLET etn h oe of model the testing MODELTEST: 1998. Crandall. A. K. and D. Posada, phy- Molecular 2007. Pryer. M. K. and Schneider, H. S., N. Nagalingum, elne,D .1965. B. D. Lellinger, of monophyly the K Investigating u o 2007. ,L .Y . B. ,F .W E. .L i R. ,W inference .L Bayesian .CKirkpatrick, MRBAYES: h 2001. i Ronquist. o F. u the and ,a Deciphering P. n 2009. J. dC .NHuelsenbeck, Pryer. .W M. a K. n g and .2 Windham, 0 D. 1 1 M. .F L., i A. r sGrusz, (Pteridaceae: ti ferns n s Cheilanthoid i g 1998. h Rollo. t s R. D. and J. G. Gastony, Gastony,G.J.andD.R.Rollo.1995.Phylogenyandgenericcircum- atn,G .adG asivc.19.Mtra neiac fthe of inheritance Maternal 1992. Yatskievych. G. and J. G. Gastony, Mu 2004. Smith. R. A. and 1988. T. J. Beitel. Mickel, M. J. and T. J. Mickel, ikPrz .A,L .Wto,adR .Hce.21.Rdfnto of Redefinition 2011. Hickey. J. R. and Watson, E. L. A., M. Link-Perez, Link,H.F.1833. i .W,L .Ko .J ohes .Eiaa .L ho,M D. M. Chiou, L. W. Ebihara, A. Rothfels, J. C. Kuo, Y. L. W., F. Li, ¨ lr . .Mu K. J., ller, li vlto ntefr genus fern the in poly- evolution interpreting ploid and delimitation species to approach diploids-first dati ferns adiantoid .Shedr 01 vdnefrrdain fcelnhi en in ferns cheilanthoid Region. of Floristic Cape radiations Greater for the Evidence 2011. Schneider. H. euneo xatfmle n eeao yohtsadferns. and lycophytes of genera and families Phytotaxa extant of sequence Botany N substitution. DNA hetero- the in genus evolution fern sporous morphological and relationships logenetic http://www.phyde.de/. from Available editor. data Phylogenetic Peiaee ntecneto hlgntcaayi of analysis phylogenetic 556–566. 59: a fern of into context the ferns. cheilanthoid in (Pteridaceae) trees. phylogenetic of the (Pteridaceae). in polyploids apomictic of origins lines. generic adjacent of and reassessment States Aliso phylogenetic United molecular southwestern Mexico—a the in Cheilanthoideae) 341–360. 85: citoso hiatodfrs( ferns from inferred cheilanthoid of scriptions hools n iohnra eoe ncelnhi ferns. cheilanthoid in genomes Botany of mitochondrial Journal American and chloroplast okBtnclGre Press. Garden Botanical York Press. Garden Botanical York New Bronx: ography. Adiantopsis Reimer. G. pa h oeDAbroefrferns. for barcode DNA core the as up ida,adK .Pyr 2011. Pryer. M. K. and Windham, 7 131–144. 17: 5 223–234. 35: Taxon 9 7–54. 19: matK ¨ yeseie tP hc sciia oortaxonomic our to critical is which P, at specimen type Fe lr .Nihi,adD unt 00 PhyDE 2010. Quandt. D. and Neinhuis, C. ller, ´ h .tei.AnAbr nvriyo Michigan. of University Arbor: Ann thesis. D. Ph. . Peiaee:Sseais iesfcto,adbioge- and diversification, Systematics, (Pteridaceae): e otsRgu oaiu Berolinensis Botanicus Regius Hortus mrcnJunlo Botany of Journal American rbcL 0 1255–1268. 60: ne Filicum Index phylogeny. osLtoiadDvdSofr;teravc and advice their Swofford; David and Lutzoni ¸ois uniaiesuyo eei eiiaini the in delimitation generic of study quantitative A Bioinformatics ytmtcBotany Systematic Marsilea uloiesequences. nucleotide Bioinformatics ´ ieaueCited Literature eBe 9 716–722. 79: ´ il (de lisle . ytmtcBotany Systematic oeua hlgntc n Evolution and Phylogenetics Molecular oehgn .Hagerup. H. Copenhagen: . h trdpye fMexico of pteridophytes The trdpyefoao aaa Mexico Oaxaca, of flora Pteridophyte 4 817–818. 14: Gaga ´ Astrolepis Taxon die rbcL 7 754–755. 17: trdca:Cheilanthoideae) Pteridaceae: 2 504–518. 32: ´ hiate yavapensis Cheilanthes a LSONE PLoS ` trnG-R pcmn rmMxc and Mexico from specimens ame la and 6 1636–1645. 96: 0 1269–1283. 60: aagermanotta Gaga (Pteridaceae). ´ matK mrcnFr Journal Fern American or eJose de moire n oteherbarium the to and ; 2 16–25. 32: :e26597. 6: antothumbs two earn eoii Apud Berolini: . rn:New Bronx: . oLayne to ; W ´ Systematic eBe complex v0.9971. Pellaea ´ gapCp lisle). . Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. nme fcoe) vrg pr imtrin diameter spore average clones); of (number wcl .J 2006. J. D. Zwickl, 859 Health, of Institutes National FERNS 1979. v1.45. CHEILANTHOID T. OF ImageJ GENUS Reeves, from NEW 2011. A Available GAGA, S. v1.5. AL.: ET Tracer LI W. 2009. Rasband, Drummond. J. A. and for A. environment Rambaut, and language A R: 2007. 2008. Salatino. Team. F. Core L. Development M. R and Salatino, A. Rodrigues, N. D. C. J., Prado, 2012] N478 X143 X150 X151 X152(4;6.324;25. 63.03±2.46; (14); JX313552 angustifolia JX313551, Mexico, Gaga JX313550, JX313453, Jalisco, JN647778, 6541, Windham 31. 70.11±2.21; NA; JX313452, Mexico, Sonora, available. not = NA sporangium. per counted spores of number tion; olcinnme vuhrlocation); accession, GenBank (voucher number and collection collector site, collection (fernlab.biology.duke.edu), number accession M. K. and Gastony, J. G. Grusz, L. A. Windham, D. M. J., C. Rothfels, cutpl,E,H cnie,L ue,M .Wnhm n .M Pryer. M. K. and Windham, D. M. Huiet, L. Schneider, H. E., Schuettpelz, cutpl,E n .M re.20.Fr hlgn nerdfo 400 from inferred phylogeny Fern 2007. Pryer. M. K. Y. and L. E. Larsson, Schuettpelz, A. Kato, M. Schuettpelz, E. Sundue, A. M. J., C. Rothfels, mt,A .18.Peiohts p –7 in 1–370 Pp. Pteridophytes. 1981. R. Pryer. A. M. Smith, K. and Yatskievych, G. Huiet, L. Windham, D. M. M., E. The Sigel, 2008. Pryer. M. K. and Windham, D. M. Grusz, L. A. E., Schuettpelz, tle .G 91 en n enale fGaeaa atII. Part Guatemala. of allies fern and Ferns 1981. G. R. G. P. and Stolze, Schneider, H. Korall, P. Schuettpelz, E. Pryer, M. K. R., A. Smith, ro,R .adA .Tyn 1982. Tryon. F. parsimony A. and using M. analysis R. Phylogenetic Tryon, PAUP* 2002. L. D. Swofford, ida,M .adE .Rb.1993. Rabe. W. E. and D. M. Windham, hn,G . .C hn,Z .Ce,H .Lu n .L ag 2007. Yang. L. W. and Liu, M. H. Chen, D. Z. Zhang, C. X. M., G. Zhang, eiyr,J .adH cnie.21.Tenwfr genus fern new The 2010. Schneider. H. and C. J. Yesilyurt, 1995. Moran. C. R. and G. Yatskievych, J. Rothfels, C. Grusz, L. A. Schuettpelz, E. Huiet, of L. D., studies M. Chromosome Windham, 2003. Yatskievych. G. and D. M. Windham, Ingroup Appendix flrebooia eunedtst ne h aiu ieiodcrite- likelihood maximum the under datasets rion sequence biological large of Physapteris A. S. U. Maryland, Bethesda, http://tree.bio.ed.ac.uk/software/tracer/. Statistical for Foundation http://www.R-project.org. R URL Computing. Austria: Vienna, computing. statistical Brazilian including from inferred Pteridaceae, species, among relationships Phylogenetic re.20.Twr monophyletic a Toward 2008. Pryer. oeua hlgntc n Evolution and Phylogenetics Molecular genera. unsampled previously of affinities Assessing the Pteridaceae: and relationships family overall fern the of phylogeny molecular A 2007. etsoagaeseisadtrepatdgenes. plastid three and species leptosporangiate II eupolypod for classification revised Polypodiales). A (Polypodiidae: 2012. ferns Pryer. M. K. and Kuo, xeric-adapted in convergence evolutionary ferns. of patterns resolving 01 pce eainhp n aiaeouini h cheilanthoid the in evolution genus farina fern and relationships Species 2011. Botany atic nuclear of utility Polypodiaceae. ferns. extant for classification A 2006. Wolf. Sciences. of Academy California Francisco: San Breedlove. eeec otoia America tropical to reference Associates. Sinauer Sunderland: 4.0. v. methods). other (*and is nihsi h hlgn fAincelnhi en ae on based ferns markers. cheilanthoid chloroplast Asian two of of sequences phylogeny the in insights First (Pteridaceae). isuiBtnclGre Press. Garden Botanical Missouri Mesoamericana and plastid ferns. Using cheilanthoid demys- 128–132. and in 99: 2009. boundaries complexes Pryer. generic species redraw M. tify to K. sequences DNA and nuclear Yatskievych, G. Beck, western Mexico. the and from States United Cheilanthoideae) (Pteridaceae: ferns cheilanthoid Press. University Oxford York: New Committee. Editorial America America North h .tei.Asi:Uiest fTexas. of University Austin: thesis. D. Ph. . — Taxon aaangustifolia Gaga .Smln nomto.Txnnm enLbDatabase Lab Fern name Taxon information. Sampling 1. Argyrochosma 3 2–629. 621– 33: (Adiantaceae) 7 712–724. 57: ,vol.2, oorp ftefr genus fern the of monograph A Knh a-e i&Wnhm65,Jlso Mexico, Jalisco, 6550, Windham & Li Fay-Wei (Kunth) a eedr97-1244 Devender van eei loih prahsfrtepyoeei analysis phylogenetic the for approaches algorithm Genetic Phytotaxa d.S .Sua .Dvde n .Kap t Louis: St. Knapp. S. and Davidse, G. Sousa, M. S. eds. , trnG-R ilin Botany Fieldiana gapCp trdpye n Gymnosperms and Pteridophytes eBn accession, GenBank rbcL h .tei.Tme rzn tt University. State Arizona Tempe: thesis. D. Ph. . :52–59. 7: nrsligplpodfr origins. fern polyploid resolving in (Pteridaceae). e ok Springer-Verlag. York: New . aaangustifolia Gaga mrcnJunlo Botany of Journal American sequences. Knh a-e i&Wnhm4989, Windham & Li Fay-Wei (Kunth) :1–522. 6: ohes3109 Rothfels en n lidpat ihspecial with plants allied and Ferns Cheilanthes 4 1172–1185. 44: Taxon Cheilanthes matK Taxon ytmtcBotany Systematic Taxon M) N479 JN647777, JN647719, (MO); 1 515–533. 61: gapCp m eBn accession, GenBank Notholaena lr fChiapas of Flora Taxon n tnaddevia- standard one ± m 6 355–368. 56: 6 369–378. 56: p 5–6 in 152–169 Pp. . Knh a-e i& Li Fay-Wei (Kunth) mrcnFr Journal Fern American Cheilanthes p 2–2 in 121–128 Pp. . DK) JN647720, (DUKE); Taxon eBn accessions GenBank 5 705–731. 55: d lr fNorth of Flora ed. , 0 1788–1800. 90: Calciphilopteris 6 1037–1050. 56: (Pteridaceae): 6 554–564. 36: d .E. D. ed. , subgenus System- lr of Flora Flora rbcL N483 X142 X153 X154 X155(4;7.532;NA. 74.75±3.26; (14); JX313585 JX313584, JX313583, JX313492, JN647813, X143 A 89±.7 NA. 68.94±4.07; NA; JX313463, ida 13 a Jose San 5133, Windham & NA. Jose NA; San JX313481, 5581, JN647803, JN647745, Windham & Li Fay-Wei Leo 90 Michoaca 4990, 64.36±1.85; (9); JX313565 JX313564, Mexico, 29. JX313461, 31. Sinaloa, JN647786, 7821, JN647728, 70.20±3.18; Windham (NY); & NA; Li Mexico, Fay-Wei JX313460, Jalisco, (Maxon) JN647785, 7820, Windham JN647727, & Li NA. (12); NA; JX313563 JX313562, NA, JX313561, NA, A., NA, S. 0301A; U. Arizona, Windham A., 32. S. 70.76±2.01; U. Arizona, NA, NA. Windham NA; JX313507, JX313540, Mexico, Tamaulipas, 64. 8090, 45.16±1.31; Windham (11); JX313560 X140 X155 X156(1;6.130;64. 60.41±3.00; (11); JX313576 JX313575, JX313480, X169 X159 X155 X154 X155() 49.26±2.16/68.36±4.85; (8); JX313595 60/27. JX313594, JX313505, Mexico, JX313539, JX313639, Tamaulipas, 7876, 21. Windham 72.23±6.74; NA; Mexico, JX313457, Chiapas, 8003, 31. 64.39±2.48; (8); 30. 17184 64.15±2.77; McVaugh (11); Mexico, JX313557 angustifolia JX313556, Sinaloa, Gaga JX313455, 7840, JN647780, Windham JN647722, & Li Fay-Wei 57/32. 40.45±4.14/58.40±2.74; (12); JX313555 3116 Rothfels eei,CsaRica, Costa Heredia, ida 02 haa,Mexico, Chiapas, 8002, Windham A .S. a-e i&Wnhm71,Cips Mexico, Chiapas, 7819, Windham & Li 32. Fay-Wei 55870 65.78±3.13; Sm.) (12); R. (A. JX313568 Mexico, JX313466, Sinaloa, JN647791, 7843, Windham Mexico, Colima, NA. 7842, Windham (16); & Li 16108 Fay-Wei NA. McVaugh Mexico, Galeotti) NA; & Jalisco, Martens JX313464, 6542, (M. JN647789, Windham JN647731, & (DUKE); Li Fay-Wei Galeotti) DK) X166 X156 X152 X167 X168 JX313619 JX313618, JX313617, 25. JX313522, 72.99±2.72; Jose (12); JX313546, 47. San JX313646, 5577, 47.16±2.41; Windham (DUKE); & (9); Li JX313574 Fay-Wei JX313479, JN647801, 4040a complanata N477 X145 X151 X152(2;6.627;26. NA. 68.16±2.72; (12); Mexico, Oaxaca, JX313572 JX313571, decomposita Mexico, JX313475, Nayarit, JN647797, 6600, Windham NA. NA; Mexico, Moraza Nayarit, Francisco 8100, decomposita Windham 53 & Montoya Li 32. 60.90±2.42; Fay-Wei NA; (Link) JX313473, Mexico, JN647795, Oaxaca, JN647737, 7837, (NY); Windham & 32. Li 64.20±3.04; Fay-Wei (12); Mexico, JX313570 JX313472, Oaxaca, JN647794, 29. 6680, 61.25±2.23; NA; Windham NA, NA, NA, a-e i&WnhmN,Oxc,Mexico, Oaxaca, NA, Windham & Li Fay-Wei A .S. a-e i&Wnhm80,Cips Mexico, Chiapas, 8000, Windham & Li Fay-Wei 40335 Sm.) R. (A. 89 aai,Mexico, Nayarit, 7839, X165 X155 X141 X159() NA. (9); JX313569 JX313471, Mexico, JX313535, Guerrero, 8103, JX313635, Windham of & Honduras, isotype Li (NY, Lempira, Fay-Wei Sm.) 8095, R. Windham (A. & Li Fay-Wei 5638 Sm.) R. (A. X147 A NA. NA; JX313477, X153() 73±.4 61. 47.31±1.74; (9); Salvador, JX313573 El 7816, ´ aachaerophylla Gaga ,Mexico, n, X164 X154 X140 A 09±.9 64. 50.99±2.39; NA; JX313470, JX313534, JX313634, ; aadecomposita Gaga DK) X163 X153 X148 A NA. NA; JX313468, JX313533, JX313633, (DUKE); CS;J673,J679,J336,N;NA. NA; JX313469, JN647793, JN647735, (CAS); N) N474 N472 X147 A 64±.7 64. 46.49±1.77; NA; JX313467, JN647792, JN647734, (NY); aaapiacea Gaga aadecomposita Gaga aachaerophylla Gaga A .S. a-e i&Wnhm72,Guatemala, 7824, Windham & Li Fay-Wei Sm.) R. (A. DK) N471 N479 X144 X153 JX313554, JX313553, JX313454, JN647779, JN647721, (DUKE); M atn aeti a-e i&Wnhm7838, Windham & Li Fay-Wei Galeotti) & Martens (M. 6555, Windham & Li Fay-Wei Galeotti) & Martens (M. M) X162 X152 X148 A NA. NA; JX313458, JX313532, JX313632, (MO); ´ itne l 23441 al. et Hinton aaarizonica Gaga ,Mexico, n, N) N473 N471 X146 X158 JX313559 JX313558, JX313456, JN647781, JN647723, (NY); N) N472 N470 X145 X156 JX313567 JX313566, JX313465, JN647790, JN647732, (NY); hiate complanata Cheilanthes ikl6064 Mickel 3119A Rothfels cutpl 461 Schuettpelz Knh a-e i&Wnhm74,Jlso Mexico, Jalisco, 7841, Windham & Li Fay-Wei (Kunth) aaangustifolia Gaga aaharrisii Gaga elr1209 Seiler aadecurrens Gaga Mce)FyWiL ida 89 Nuevo 7879, Windham & Li Fay-Wei (Mickel) M atn aeti a-e i&Windham & Li Fay-Wei Galeotti) & Martens (M. M atn aeti a-e i&Windham & Li Fay-Wei Galeotti) & Martens (M. ohes2689 Rothfels ´ relv 20549 Breedlove cag 16427 McVaugh ot Rica, Costa , timn 4721 Steinmann N) N470 N478 X146 NA; JX313476, JN647798, JN647740, (NY); aadecurrens Gaga M atn aeti a-e i& Li Fay-Wei Galeotti) & Martens (M. M atn aeti a-e i& Li Fay-Wei Galeotti) & Martens (M. aaarizonica Gaga Mxn a-e i&Wnhm3168, Windham & Li Fay-Wei (Maxon) aaapiacea Gaga DK) N478 N476 JX313474, JN647796, JN647738, (DUKE); DK) N476 N474 EU268673, JN647784, JN647726, (DUKE); aaapiacea Gaga N) N472 N480 JX313478, JN647800, JN647742, (NY); Mxn a-e i&Wnhm5574, Windham & Li Fay-Wei (Maxon) TX;J672,J678,JX313459, JN647783, JN647725, (TEX); aachaerophylla Gaga aacuneata Gaga atooe 2516 Bartholomew ohes3288 Rothfels relv 55788 Breedlove ohes3183 Rothfels Mce)FyWiL Windham & Li Fay-Wei (Mickel) ig n e 2378 Nee and Diggs ohes08-023 Rothfels ´ Knh a-e i&Windham & Li Fay-Wei (Kunth) .R m var. Sm. R. A. aaarizonica Gaga ´ ot Rica, Costa , ot Rica, Costa , DK) N474 JN647802, JN647744, (DUKE); ese n ese 18 Webster and Webster aahirsuta Gaga ohadFyel89200 Fryxell and Koch CS;J672,JN647782, JN647724, (CAS); N) N471 JN647799, JN647741, (NY); M) N470 JN647788, JN647730, (MO); aaangustifolia Gaga ida ta.566 al. et Windham Mce)FyWiL & Li Fay-Wei (Mickel) Mce)FyWiL & Li Fay-Wei (Mickel) Mce)FyWiL & Li Fay-Wei (Mickel) ikladLoad4881 Leonard and Mickel Mxn a-e i& Li Fay-Wei (Maxon) aaharrisii Gaga nesn12567 Anderson aacuneata Gaga Ln)FyWiL & Li Fay-Wei (Link) aacuneata Gaga DK) JN647736, (DUKE); DK) JN647739, (DUKE); aachaerophylla Gaga rs 105 Grusz DK) JN647755, (DUKE); aacomplanata Gaga ohese l 2618 al. et Rothfels aagermanotta Gaga interrupta CS;JN647743, (CAS); aacomplanata Gaga aacomplanata Gaga Ln)FyWiLi Fay-Wei (Link) aacomplanata Gaga Mxn Fay-Wei (Maxon) N) JN647733, (NY); M atn & Martens (M. aaarizonica Gaga relv 18570 Breedlove ´ aacuneata Gaga ,Honduras, n, F;JX313640, (F); ohes3110 Rothfels iln149 Millan Breedlove Breedlove (DUKE); (DUKE); (Maxon) Mickel); (Kunth) (TEX); Moran (Link) (Link) (NY); (NY); Gaga Gaga Gaga Lloyd Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Tue, 23 Oct 2012 13:19:45 Copyright (c) American Society for Plant Taxonomists. All rights reserved. N488 X146 A NA. NA; JX313486, JN647808, Mexico, Morelos, 24. 65.05±1.97; DK) N472 N480 X148 A NA. NA; Mexico, JX313488, Mexico, JN647810, 6462, 32. Windham JN647752, 63.02±3.38; & (DUKE); Li (12); Fay-Wei JX313579 (Kunze) JX313487, JN647809, Mexico, 61. 49.35±1.70; (14); JX313577 ida 12 a Jose San 5132, Windham A NA. NA; 53 Leo 49.41±1.43; (8); JX313578 hintoniorum Leo JX313483, Nuevo JN647805, 7877, JN647747, (TEX); Windham & Li Fay-Wei 32. 62.37±2.22; (16); Leo JX313623 JX313622, hintoniorum 6837 JX313524, al. JN647834, et (14); Hinton JX313616 NA. JX313615, JX313614, Mexico, JX313520, Mexico, 29. 69.05±2.67; Chiapas, JN647832, 7830, Windham JN647774, & Li Fay-Wei N481 X159 X162 X163() NA. (7); JX313613 Mexico, JX313612, Jalisco, JX313519, JN647831, 6624, Windham & X158 X160 X161(2;NA. (12); JX313611 Mexico, JX313610, JX313518, Jalisco, NA. 6583, NA; Mexico, JX313517, Hidalgo, 6490, NA. (18); JX313609 JX313608, Mexico, NA. (11); 2696 Rothfels N470 N488 X147 A NA. NA; JX313497, JN647818, JN647760, NA. (12); JX313588 2839 Diez JX313496, JN647817, marginata Ecuador, NA. Pichincha, JN647759, 7443, (14); Windham (DUKE); & JX313587 Li Fay-Wei JX313495, (Kunth) JN647816, JN647758, Rica, NA. Costa Heredia, 5579, NA; 2688 Windham & JX313494, Li Fay-Wei JN647815, (Kunth) Rica, Costa JN647757, 32. Rica, Heredia, 5571, 69.24±2.92; (DUKE); Windham Costa & NA; Li Heredia, Fay-Wei JX313493, (Kunth) 5569, JN647814, Windham JN647756, & Li 32. 64.59±1.70; (9); JX313582 Bolivia, JX313491, Tarija, 4986, NA. (13); Moraza JX313581 Francisco 6238b 8099, Mickel Windham & lerstenii Li Fay-Wei Honduras, Beitel) & (Mickel 109 Kne a-e i&Wnhm68,SnLi Potosı 37 Luis [Volume San 6985, Windham & Li Fay-Wei (Kunze) BOTANY SYSTEMATIC hirsuta Gaga 860 Knh a-e i&Wnhm73,Pel,Mexico, Puebla, 7833, Windham & Li Fay-Wei (Kunth) Mexico, Jalisco, 6571, Windham & Li Fay-Wei ´ ´ ,Mexico, n, Mexico, n, DK) N473 N481 X149 A NA. NA; JX313489, JN647811, JN647753, (DUKE); aahirsuta Gaga DK) A A A X156(0;NA. (10); JX313586 NA, NA, NA, (DUKE); ikladHlwg3891 Hellwig and Mickel ida ta.519 al. et Windham ohes3023 Rothfels Mce etl a-e i&Wnhm83,Oxc,Mexico, Oaxaca, 8237, Windham & Li Fay-Wei Beitel) & (Mickel aakaulfussii Gaga N) N479 N477 X142 A NA. NA; JX313462, JN647787, JN647729, (NY); aahirsuta Gaga Knh a-e i&Wnhm72,Mxc,Mexico, Mexico, 7823, Windham & Li Fay-Wei (Kunth) adrn066 Calderon DK) N479 N487 X155 X165 JX313606 JX313605, JX313515, JN647827, JN647769, (DUKE); Mneh eo)FyWiL ida 88 Nuevo 7878, Windham & Li Fay-Wei Nesom) & (Mendenh. Nuevo 7813, Windham & Li Fay-Wei Nesom) & (Mendenh. Ln)FyWiL ida 52 a Jose San 5582, Windham & Li Fay-Wei (Link) U,ioye;J333,J333,J339,JX313580, JX313490, JX313536, JX313636, isotype); (UC, aahirsuta Gaga itne l28587 al et Hinton itne l22695 al et Hinton aakaulfussii Gaga Ln)FyWiL ida 80 eio Mexico, Mexico, 7880, Windham & Li Fay-Wei (Link) asivc n atn 89-286 Gastony and Yatskievych aamarginata Gaga L,ioyeof isotype (LL, DK) N470 N488 X156 JX313607, JX313516, JN647828, JN647770, (DUKE); aahirsuta Gaga ern ta.6089 al. et Serrano ´ M) A A A A 10±.5 63. 51.09±1.65; NA; NA, NA, NA, (MO); Ln)FyWiL ida 42 Hidalgo, 6482, Windham & Li Fay-Wei (Link) ot Rica, Costa , ohes3154 Rothfels Kne a-e i&Wnhm40,Morelos, 4407, Windham & Li Fay-Wei (Kunze) ohes3032 Rothfels DK) N479 N487 X145 NA; JX313485, JN647807, JN647749, (DUKE); Ln)FyWiL ida 82 Oaxaca, 7832, Windham & Li Fay-Wei (Link) aahirsuta Gaga aahintoniorum Gaga N) N475 N483 X151 NA; JX313521, JN647833, JN647775, (NY); aakaulfussii Gaga Kne a-e i&Wnhm5073, Windham & Li Fay-Wei (Kunze) TX;J674,J670,JX313484, JN647806, JN647748, (TEX); N) N476 N484 JX313482, JN647804, JN647746, (NY); ohes08-022 Rothfels Ln)FyWiL Windham & Li Fay-Wei (Link) ohes3212 Rothfels Knh a-e i&Windham & Li Fay-Wei (Kunth) aamarginata Gaga hiate spiculata Cheilanthes aahirsuta Gaga DK) N472 JN647830, JN647772, (DUKE); DK) N471 JN647829, JN647771, (DUKE); ´ Ln)FyWiL Windham & Li Fay-Wei (Link) ,Mexico, n, M) N474 JN647812, JN647754, (MO); aamarginata Gaga aamarginata Gaga ohes2693 Rothfels Kne a-e i& Li Fay-Wei (Kunze) ohes3137 Rothfels Mneh Nesom) & (Mendenh. relv 22395 Breedlove aahirsuta Gaga DK) JN647773, (DUKE); DK) JN647751, (DUKE); etr 1393 Ventura itne al et Hinton Ln)FyWiLi Fay-Wei (Link) ´ M) JN647750, (MO); aamarginata Gaga aamarginata Gaga aamarginata Gaga Mexico, , aakaulfussii Gaga aakaulfussii Gaga Knh Fay-Wei (Kunth) aamarginata Gaga aalerstenii Gaga ´ ohes2988 Rothfels ;JN647776, ); ohes3731 Rothfels ohese al. et Rothfels 2692 Rothfels ot Rica, Costa , (DUKE); (DUKE); (Kunth) (Kunth) Larsson Tejero- (Link) 20157 (NY); (NY); Gaga Gaga Gaga Gaga ´ n, a-e i&Wnhm73,Oxc,Mexico, Oaxaca, NA. 7834, NA; Windham JX313498, JN647819, & JN647761, Li Fay-Wei eiaie l 1363 rufopunctata Cheilanthes al. et Deginani NA. 89-219 Gastony NA; and Yatskievych A., 62. S. U. A., 58. S. 2.161; 52.004± U. NA; California, NA, Lellinger NA. (9); A., S. U. A., 64. S. 37.602±1.548; U. California, NA. (11); JX313548 A., S. U. California, e i&Wnhm71,Gerr,Mexico, Guerrero, 7815, Windham & Li NA. Wei (13); JX313621 JX313620, JX313523, N472 N480 X149 A NA. NA; Mexico, JX313499, Jalisco, JN647820, 7844, JN647762, Windham & Li Wei X168 X158 X153 A NA. NA; JX313503, JX313538, JX313638, 59. 49.88±3.91; (10); JX313596 Mexico, 60. 59767 Breedlove pellaeopsis 4394 ida 14 eio Mexico, Mexico, 7154, Windham 64. 43.33±1.22; (15); JX313593 JX313592, e i&Wnhm89,Crao ot Rica, Costa Cartago, 8096, Windham & Li Wei 23±.7 59. 42.37±1.97; Potosı Luis San 8236, Windham 4881 63. & 49.46±2.53; Li 60. Fay-Wei NA; Reeves) (T. JX313513, 43.51±1.58; JX313544, Quere JX313644, 8093, Windham (16); (NY); & Li Fay-Wei JX313603 Reeves) (T. JX313602, JX313512, Mexico, Guanajuato, 60. 45.48±1.20; 62. 46.75±2.25; 586 (17); Kenoyer JX313599 NA. JX313510, purpusii Mexico, Gaga JN647826, Hidalgo, (13); 6486, JN647768, Windham & (DUKE); JX313598 Li Fay-Wei JX313597, Reeves) (T. JX313509, JN647825, Quere 4995, Windham 86 aaa Mexico, Oaxaca, 32. 7846, 66.03±2.13; 64. 48.22±2.23; (9); 7431 JX313589 Mickel JX313500, membranacea JX313537, JX313637, Br.f)T or 38 oihma,Laos, NA. Bolikhamsai, 5318, NA; Moore JX313448, T. f.) JX313528, (Burm. JX313628, (US); X169 X159 X149 A NA. NA; JX313449, JX313529, JX313629, ae 71 rna Tanzania, Iringa, 4711, Baker .Wlew 89 rgn .S A., S. U. Oregon, 3869, Wollenw. E. NA. NA; JX313450, X151 X141 A NA. NA; JX313451, JX313531, Outgroup sioi meifolia Aspidotis aapellaeopsis Gaga N) N475 N483 X154 A 56±.0 62. 45.62±1.60; NA; JX313504, JN647823, JN647765, (NY); U,ioye;J334,J334,J331,J330 (9); JX313604 JX313514, JX313545, JX313645, isotype); (UC, li ta 29543 al et Iltis re 06-02 Pryer sioi carlotta-halliae Aspidotis ida 823 Windham hiate micropteris Cheilanthes N) N473 N481 X151 X150 X151(9); JX313591 JX313590, JX313501, JN647821, JN647763, (NY); — Mce)FyWiL ida 01 aaa Mexico, Oaxaca, 8001, Windham & Li Fay-Wei (Mickel) F;J334,J334,J331,J330,J330 (14); JX313601 JX313600, JX313511, JX313542, JX313642, (F); Dvn. a-e i&Wnhm74,Oxc,Mexico, Oaxaca, 7845, Windham & Li Fay-Wei (Davenp.) T evs a-e i&Wnhm88,Hdlo Mexico, Hidalgo, 8089, Windham & Li Fay-Wei Reeves) (T. sioi californica Aspidotis Gaga aapurpusii Gaga aamembranacea Gaga CS;J676,J672,J330,N;47.74±1.68; NA; JX313506, JN647824, JN647766, (CAS); etgam triangularis Pentagramma sioi densa Aspidotis sioi californica Aspidotis eza 178 Metzgar sp. etr n oe 9059 Lopez and Ventura sioi densa Aspidotis M) X167 F515 U663 A NA. NA; EU268683, EF452145, JX313627, (MO); DK) X165 X156 X146 JX313549 JX313446, JX313526, JX313625, (DUKE); D .Etn i.Sr.55,Tmuia,Mexico, Tamaulipas, 5056, Serm. Pic. Eaton) C. (D. ´ Mce)FyWiL ida 14 Jalisco, 8104, Windham & Li Fay-Wei (Mickel) ikl7057 Mickel ao Mexico, taro, ida 856 Windham 07 a Jose San 8097, N,ioye;J334,J334,JX313508, JX313541, JX313641, isotype); (NY, DK) A A A A 163 2.613; 41.663± NA; NA, NA, NA, (DUKE); oes.72,Crac,Bolivia, Carrasco, 7829, Rosenst. T evs a-e i&Wnhm8092, Windham & Li Fay-Wei Reeves) (T. aapurpusii Gaga aob 2654 Kayombo DK) X164 X155 JX313445, JX313525, JX313624, (DUKE); ek1229 Beck M) X166 X157 JX313447, JX313527, JX313626, (MO); N) N474 N482 JX313502, JN647822, JN647764, (NY); akr902 Ranker Dvn. a-e i&Windham & Li Fay-Wei (Davenp.) Bak)Lligr37,Oregon, 3870, Lellinger (Brack.) ´ Ho. ut xCpl 3831, Copel. ex Nutt. (Hook.) Bak)LligrN,Washington, NA, Lellinger (Brack.) timn 2528 Steinmann w 79 iins Argentina, Misiones, 3709, Sw. aamonstraparva Gaga ot Rica, Costa , WH anr&EF Gilbert) E.F. & Wagner (W.H. ele angulosa Pellaea DK) A A A NA; NA, NA, NA, (DUKE); re 06-01 Pryer aapellaeopsis Gaga aamarginata Gaga aamarginata Gaga Ho. ut xCpl NA, Copel. ex Nutt. (Hook.) otnadBad 2306 Brandt and Boutin DK) X167 JX313547, JX313647, (DUKE); Kuf)Ytk,Wnhm& Windham Yatsk., (Kaulf.) M) X160 JX313530, JX313630, (MO); N) X163 JX313543, JX313643, (NY); T evs a-e i& Li Fay-Wei Reeves) (T. ´ ao Mexico, taro, ue l WS-296 al. et Wu DK) A A NA, NA, NA, (DUKE); nesnadLaskowski and Anderson ais 24959 Davidse eintsarifolia Hemionitis oa 2361 Moran ikl7432 Mickel DK) JX313631, (DUKE); M) JN647767, (MO); ´ Mexico, , Br xWilld.) ex (Bory aapurpusii Gaga aapurpusii Gaga aapurpusii Gaga Mce)Fay- (Mickel) a-e i& Li Fay-Wei Knh Fay- (Kunth) Knh Fay- (Kunth) ohes3027 Rothfels arg 6598 Barriga ude613 Sundue Purpus (MO); (MO); (MO); (NY); (NY); Gaga Gaga