Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 aooi concept. taxonomic Cheilanthes and 2012), rusto species several groups of transfers recent by somewhat improved al. been every has situation nearly et The Cheilanthoideae. subfamily in the found Yesilyurt of genus clade 2011; major this to 1995, al. assigned Rollo species et with and Eiserhardt 2015), Gastony 2007; adequate (e.g. Kirkpatrick an ferns 1998; with cheilanthoid study of molecular sampling every by polyphyletic be brandegeei Cheilanthes of better 2014). transfer recent signature the mor- senae (e.g. genetic of Adiantum another in degree whose inclusion some their but to supports due similarity, genus one phological within have that placed species enigmatic been several pinpointed also have data aeo ulcto uut6 2019 6, August publication Taxonomists of Plant Date of Society American 10.1600/036364419X15620113920536 the DOI by 2019 Copyright © Botany Systematic Yslute l 2015), al. 2012), et al. (Yesilyurt et (Li Windham & 2010), Schneider and (Yesilyurt H.Schneid. e.g. genera; F´ recircumscribed or new been as have segregated that subgroups identi- monophyletic supported diversification, well cheilanthoid fying phylogenetic cheilanthoids reliable studying a for of established framework have 2011; 2015) sampling al. al. et et Eiserhardt broader 2009; Yesilyurt al. a et Windham 2007; of (Kirkpatrick analyses in- lecular ferns, cheilanthoid of and genera large con- cluding 1998) 1982). several 1995, 1973, that Rollo Tryon and firmed (Gastony and studies (Tryon molecular evo- environments Early convergent arid extensive in to these lution attributed and often alone, morphology dif- are on notoriously difficulties based been habitats circumscribe have to seasonally-xeric group ficult to this in adapted Genera species worldwide. 400 including family, than the half more nearly comprises 2016) I Cheilanthoideae PPG 2007). (sensu al. et (Schuettpelz families fern diverse 03,and 2013), e(Link-P´ ee h ou forsuyi n uhseis urnl nw as known currently species, such one is study our of focus The trdca soeo h otseisrc n ecologically and species-rich most the of one is Pteridaceae eas fisdsiciemrhlg ihntebmeidcae(int efacietr,wl-eeoe suonuim n narrow, and pseudoindusium, well-developed architecture, all leaf to (pinnate sister is clade genus it new bommeriid that the propose evidence the we and here within ferns, scales), cheilanthoid morphology rhizome other red-brown distinctive all concolorous to its clade of sister the Because bommeriids, the of member el xaddbmeidcae efidmn faiua rcoe,rtclt-rsaeprsoemrhlg,adltrliiito fthe of initiation lateral and morphology, perispore reticulate-cristate trichomes, acicular of indument unit leaf meristem. that characters clade: gametophyte three identify bommeriid We cheilanthoids. America expanded across South homoplasious in newly extensively congeners are hypothesized a petioles) with as fractiferous than species and rather this pseudoindusium, Mexico, classify developed mainland to and used Peninsula traditionally characters California Morphological Baja Africa. the and in distribution a have that taxa other he lsi akr ( markers plastid three iagog@ueeu re@ueeu z.ogkodk.d;lyehitdk.d;[email protected] [email protected]; [email protected]; [email protected]; [email protected]; Pellaea Keywords Abstract Baja Cheilanthes iaO ere ahenM Pryer, M. Kathleen George, O. Lisa Myriopteris Adiantopsis Ormopteris ean natfca n eaieyuninformative relatively and artificial an remains ik eeide oyhltc usqetmo- Subsequent polyphyletic. indeed were Link, 21) 43:p.471 pp. 44(3): (2019), — rze l 2011), al. et erez — e oopcfcGnsSgeae from Segregated Genus Monospecific New A : h hlgntcpsto of position phylogenetic The aaCalifornia, Baja ae into Baker Sw., eateto ilg,Dk nvriy uhm ot aoia27803,USA; 27708-0338, Carolina North Durham, University, Duke Biology, of Department D.C.Eaton. GuzadWnhm21) u currently but 2013), Windham and (Grusz atpA (Link-P´ .m Yslute l 05.Molecular 2015). al. et (Yesilyurt J.Sm. Myriopteris Doryopteris , – 482 rbcL Adiantopsis Bommeria rze l 2011), al. et erez Lytoneuron , trnG-R Cheilanthes Calciphilopteris F´ e(rs n Windham and (Grusz ee omri en,celnhi en,gmtpyedvlpet lsi phylogeny. plastid development, gametophyte ferns, cheilanthoid ferns, bommeriid , n oprtv opooy eew rsn outeiec o h eonto of recognition the for evidence robust present we Here morphology. comparative and ) J.Sm., hiate brandegeei Cheilanthes ySheteze al. et Schuettpelz by 1 uhrfrcrepnec ([email protected]) correspondence for Author Kozc)Yesilyurt (Klotzsch) Gaga a ensonto shown been has Notholaena omnctn dtr ljnr Vasco Alejandra Editor: Communicating Gaga re,F.W.Li Pryer, eiyr & Yesilyurt Adiantopsis 1 L tal. et (Li z-ogKo an ue,adMcalD Windham D. Michael and Huiet, Layne Kao, Tzu-Tong enedmct h aaClfri eisl fMxc,wsivsiae using investigated was Mexico, of Peninsula California Baja the to endemic fern a , R.Br., 471 Baja xii pret es eoiso elwo rneflavonoids orange or ( yellow three of deposits other dense the to sparse whereas exhibit pubescent, sparsely as characterized species, the bases. of petiole two of the leaves The on scales thin large, with petioles fractiferous ae,and Baker, taxa Mexican marily h nenendemic Andean species,the other four included group This (1982). Tryon and AmericanTryon of groups species formal oteBj aionaPnnuao eio(emn21) twasthenamesakeofthe It 2018). (Rebman Mexico of Peninsula California Baja the to Arbel´ ochracea N. pce eetaserdto transferred were species hscaehv l enasge otegenus the to assigned been all have clade this of group earliest-diverging cheilanthoids the clade, bommeriid in the cited of that data, 2004) unpub. Smith (Cranfill and Mickel Cranfill the on- by supported our ferns suggestion when cheilanthoid earlier then, of surprising, surveys somewhat molecular was going It 2009). al. (Windham clade et hemionitid large the within to nested related deeply closely considered are most species be to non-farinose 1972) the (1960, Tryon that by reveal prep.) in al. Neo- et and African between affinity pteridophytes. floristic tropical of dispersal as indicator (2001) long-distance Smith and an Moran of by cited example and speciation by an followed as once interpreted was species again species these peti- among separation fractiferous geographic African/Namibian and The oles. morphologies leaf South similar superficially the to deltoidea C. related that migration. speculated closely long-distance also had after (1960) speciation Tryon of example classic relative closest presumed of aration Arbel´ and (Yatskievych “ farina oacmoaei.Orrslsplace results Our it. accommodate to hiate brandegeei Cheilanthes eetmlclraaye Esrad ta.21;Windham 2011; al. et (Eiserhardt analyses molecular Recent sep- geographic wide the that hypothesized (1972) Tryon Cheilanthes e,and aez, ” nteraailla ufcs eety h farinose the Recently, surfaces. leaf abaxial their on ) uz and Kunze .palmeri C. hiate brandegeei Cheilanthes Ho. as.&Arbel´ & Yatsk. (Hook.) ” pten fsoaga itiuin rsneo well- a of presence distribution, sporangial of (patterns Cheilanthes Wnhme l 09.Teohrfv pce in species five other The 2009). al. et (Windham .jaliscana N. “ ..ao,adwsdfndb having by defined was and D.C.Eaton, .aurantiaca C. .fractifera C. .brandegeei C. .capensis C. e 2008). aez sasal okdeln enendemic fern rock-dwelling small, a is .fractifera C. Notholaena as.&Arbel´ & Yatsk. .brandegeei C. Bommeria .brandegeei C. .l (Pteridaceae) l. s. aabrandegeei Baja Tub)S. ae ntheir on based Sw., (Thunb.) ..ro,pu he pri- three plus R.M.Tryon, Bj aiona rmits from California) (Baja aez, group, Cheilanthes Cv)T.Moore, (Cav.) pce ihnta clade. that within species n r urnl named currently are and Prva ne)wsa was Andes) (Peruvian .brandegeei C. .aureolina N. ih eamember a be might ” and .brandegeei C. n feee in- eleven of one e,respectively aez, .fractifera C. oehrwith together eonzdby recognized .brandegeei C. Bommeria ih be might as.& Yatsk. .aurea C. ethe sa as “ core ,are Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 ae nsmlrt oprsn.Abgosyaindrgoscue by adjusted caused manually regions aligned then Ambiguously and comparisons. 2014), similarity (Larsson on 1.19 based Aliview in 2004) (Edgar ( of sporangia. the distributed submarginally protecting lobed, pseudoindusium deeply marginal a well-differentiated and blades leaf triangular pinnate-pinnatifid, curdfrti td n r eoie nGnak(pedx1). (Appendix GenBank in deposited newly were are sequences and 28 study of this total for A Sequencher acquired 2011). using Corporation assembled Codes and (Gene 5.0.1 edited manually were chromatograms ( gene trnR partial with gene partial sequenced: modifica- were markers and plastid following Three amplified (2007). Georgia), Pryer and Norcross, Schuettpelz in Bio-tek, described tions (Omega SP E.Z.N.A. kit the either or DNA California) using Valencia, plant (Qiagen, material kit mini herbarium plant and/or DNeasy the silica-dried, fresh, from isolated indicated are 1. numbers Appendix accession GenBank in and Vouchers to outgroup. group our sister as chose apparent the we H.Schneid., phylogenetic & 2011; 2015), Yesilyurt al. previous et clades al. on Eiserhardt 2007; et major Based al. et Yesilyurt the (2009). (Schuettpelz ferns of al. cheilanthoid of et five studies Windham representing by genera recognized cheilanthoid other seven sindto assigned oopcfcgns eenadhneot named henceforth Treatment). Taxonomic and herein genus, monospecific distinct of relatively the segregate to species chosen recognized We five group. Bommeria the this to within sister relationships be may that explaining confirm that gametophyte in and sporophyte useful the both of acters of taxa recognized all as well Bommeria as it, to related be to hypothesized rbcL from distinct very is superficially, least 472 20) and (2007); osdrdms lsl eae oi ( it to related capensis C. closely most considered species target our brandegeei, of individuals two including specimens, twenty-one atpA unprotected veins. and the hairs, along acicular scattered chei- sporangia with other blades pinnatifid), intricately most leaf often pentagonal (but from Haufler simple its distinguished and on based easily Ranker lanthoids is 1990; which Ranker 1990), 1979; (Haufler E.Fourn. rbcL trnG-R obnd2 4749 21/3817 6365/1. GTR 15.5 / 635 26.3 / 1076 3.8 / 3241 4094 4417 21 combined atpA marker Plastid eew netgt h upiigpyoeei placement phylogenetic surprising the investigate we Here Table hlgntcAnalysis Phylogenetic Sequencing and Amplification, Extraction, DNA ao Sampling Taxon hiate brandegeei Cheilanthes , negncsae ( spacer intergenic , ; trnG-R rbcL 10b)a ecie nSheteze l 21) N sequence DNA (2015). al. et Schuettpelz in described as bp) 1100 .Sqec hrceitc n etftsqec vlto oes utpemdl r rsne nodro oo oiinpartitions. position codon of order in presented are models Multiple models. evolution sequence best-fit and characteristics Sequence 1. , h i aata ro 16,17)adTynadTyn(1982) Tryon and Tryon and 1972) (1960, Tryon that taxa six the ,and ohleaaroia .jaliscana, N. aureolina, Notholaena eas rtclyreaiemrhlgclchar- morphological reexamine critically also We . trnG-R Bommeria Taxa 114 8122 . 8 1120/1. GTR 11.6 / 210 21.1 / 383 5.9 / 2229 1811 1847 21 110 399 . 4 8711/08GTR /10.8 141 18.7 / 245 0.4 / 97 1309 1309 21 116 7 1 . 4 6028/2. HKY 25.5 / 248 46.0 / 448 4.5 / 915 974 1261 21 samrhlgclychsv ru,w have we group, cohesive morphologically a is # na nlssta nldsalseispreviously species all includes that analysis an in ) .brandegeei C. trnG-R atpF lgmn length Alignment negncsae ihpartial with spacer intergenic — aeil n Methods and Materials yRne n afe 19) n h yeseisof species type the and (1990), Haufler and Ranker by aeilfrDAetato a bandfrom obtained was extraction DNA for Material gene, (bp) ; ,sqecswr is lge sn MUSCLE using aligned first were sequences ), 80b) sdsrbdi cutpl n Pryer and Schuettpelz in described as bp), 1830 — o aho h he lsi akrdatasets marker plastid three the of each For atpF samme ftebmeidclade, bommeriid the of member a is sn he lsi akr ( markers plastid three using - atpA acpiotrsludens Calciphilopteris rbcL nldd(bp) included Characters negncsae,adpartial and spacer, intergenic ee(39b)adcomplete and bp) (1309 gene hiate rciea .deltoidea C. fractifera, Cheilanthes and hiate brandegeei Cheilanthes .ochracea N. trnG .brandegeei C. Bommeria Bommeria — isn data Missing “ b %) / (bp eeadpartial and gene oecheilanthoids, core eoi N was DNA Genomic Wl.e Hook.) ex (Wall. ,alfv species five all ), ihfully with YTMTCBTN Vlm 44 [Volume BOTANY SYSTEMATIC Because . Baja sanew a as aibesites Variable b %) / (bp atpA atpA (see atpA trnR Baja ,at ” - , , nomtv sites Informative 8,K0 K,SM T)adfu mn-iert aitos(equal, JC, variations sets: rate model among-site four (six and 1 schemes GTR) SYM, substitution HKY, three K80, partition F81, within per model selected best then The was positions. codon by and coding/non-coding are sites informative 1. phylogenetically Table and in percentages Alignment sites, summarized as variable well analysis. data, as analyses, missing in further of included characters from of number excluded lengths, were insertions/deletions ieiodsoeaogfu elctswsslce stebs Ltree; ML best with replicates, the 1000 as from “ calculated selected was (MLBS) was ML for replicates support bootstrap four among score likelihood 1. with Table 2006), in provided are dataset each for models PAUP fitting in (BIC) criterion formation iulyisetdadcmae o ofit sn otta values bootstrap using conflicts for compared and inspected visually burn-in. as discarded the were and trees generations, 1.6 of 1000 v. every 25% sampled Tracer con- first were using Trees The confirmed 2014). generations. al. and et 1,000,000 examined (Rambaut and was parameters chains of four independent vergence two with with each 2012) al. runs, et (Ronquist MCMC 3.2.3 v. MrBayes in out carried $ rp fgyeo nagassie n aulycutn prsper spores counting manually and slide, glass a small on in glycerol sporangia of individual opening drops Treatment. specimens, herbarium Taxonomic select 1 the removing from by of conducted were section analyses These Examined Specimens Additional brandegeei Baja Bommeria brandegeei ( ameter 20 8 at at graphed stereomicroscope 12.5 MZ Leica 40 a these using of taken many were of were photographs characters morphology Representative species. pseudoindusial In all and for symmetry. petioles, characterized distribution, fractiferous gametophyte of sporangial absence of and or extent presence indument, color, scale Ranker leaf rhizome genus addition, and dissection, the (1979) delimiting and in Haufler useful shape by most We be determined to Treatment). characters (see (1990) Taxonomic those clade of on bommeriid section focused Examined recircumscribed Specimens newly Additional the across morphologies subpaleacea B. Haufler, ehrenbergiana meria Re- of Digital specimens Dryad the in 2019). deposited al. are et Phy- trees (George 1). pository phylogenetic (Table and outgroup sequences the sets new using data concatenated 28 rooted the and were respectively; trees characters logenetic bp, 4094 974 included described set and as data 1309, 2012) 1811, al. comprised et the Bayesian/ study (Ronquist of and 3.2.3 portions 2006), The v. (Zwickl 4.0a159 above. MrBayes 2.0 PAUP* in Garli in analyses in selection MCMC analyses model/partition ML best 2003), single to (Swofford a into subject combined were and they dataset datasets, incongruencies three topological the among no observed Because were 1996). Kellogg and Mason-Gamer genthreshfortopoterm b )Bs-i oe (partitioned) model Best-fit %) / (bp ro oeouinr oe eeto,ec aae a attoe by partitioned was dataset each selection, model evolutionary to Prior aiu ieiodaaye eecridotuigGri20(Zwickl 2.0 Garli using out carried were analyses likelihood Maximum retplge eeae rmec niiulpatdrgo were region plastid individual each from generated topologies Tree ubro prsprsoagu 3 s 4 n vrg pr di- spore average and 64) vs. (32 sporangium per spores of Number 0 rBysa otro probabilities posterior Bayesian or 80% opooyo prpye,Soe,adGametophytes and Spores, Sporophytes, of Morphology I, 3 1 anfcto.Dymutdsie fhisadsae eephoto- were scales and hairs of slides mounted Dry magnification. G, m .hispida B. swl sasalrnme fseiesrpeetn all representing specimens of number smaller a as well as , pce n aapeiul yohszdt ecoeyrltdto related closely be to hypothesized previously taxa and species )wr eoddfo ieenhraimseiesof specimens herbarium nineteen from recorded were m) “ 1 genthreshfortopoterm aabrandegeei Baja oce pcmn o hs pr tde r itdi the in listed are studies spore these for specimens Voucher . ao eecrflyeaie ocmaesporophytic compare to examined carefully were Maxon I 3 1 magnification. Mt.e un Underw., Kuhn) ex (Mett. oigregions) coding GTR position); oiin;HKY position); oigregions) coding GTR position); )uiganeighbor-joini a using G) Kozc)Underw., (Klotzsch) 1 1 1 1 ” frtcdnpsto) HKY position); codon (first G frtcdnpsto) K80 position); codon (first I frtcdnpsto) HKY position); codon (first I e o2,0.TeBysa/CCaaye were analyses Bayesian/MCMC The 20,000. to set G atpA DC ao)Wnhm&L.O.George, & Windham Eaton) (D.C. , rbcL ” 1 1 1 .a5 Sofr 03.Tebest The 2003). (Swofford 4.0a159 * e o10000 h rewt h best the with tree The 1,000,000. to set tidcdnpsto n non- and position codon (third G tidcdnpsto n non- and position codon (third G ,and tidcdnposition) codon (third G .elegans B. trnG-R $ .pedata B. – 9 Hli n ul1993; Bull and (Hillis 99% gte n aeinin- Bayesian and tree ng aue natsporangia intact mature, 5 acpiotrsludens Calciphilopteris Bommeria ee nlzdi this in analyzed genes Dvn. akr& Ranker (Davenp.) 1 1 S. .or. and E.Fourn., (Sw.) 1 scn codon (second I scn codon (second I scn codon (second I nldn leaf including , — Herbarium 3 Bom- .All Baja and Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 il,wl-eie rnvreasiso oe ertebase) in the ubiquitous near zones are abscission mul- transverse (with well-defined petioles tiple, Fractiferous scales. light-brown colorous, axis; central whereas most brown, of reddish those to orange distinctly and concolorous 2. Table aabrandegeei across Baja characters morphological of investigation comparative ar(S8% P10.Rltosisaogti group, this among Relationships 1.0). PP hispida, 89%, (BS pair and taxa, sister as resolved are of species Among five all to sister as brandegeei nawel ai vrapro f1 o orpae,wt healthy with presence the gametophytes. plates, whether influenced assess Four gametophytes to spores mature mo. of of sowing 12 second a of to subject period were ap- four-month-old a at of over day basis populations per weekly light a fluorescent on of h of Observations 12 23°C. proximately with up right-side p positioned These plates. petri diameter mm 60 a eoe from removed far type the to related of closely species 93%, clade notholaenid (BS the group within well-supported 1.0) a PP form 1), Fig. in respectively, upre (BS supported 2019] hra hs of those whereas brandegeei of species all in observed of dissection and shape pedata Bommeria 50 at stereomicroscope 12.5 MZ Leica a on sporangium prswr ondrcl noHevly onto directly sown were spores sec Examined Specimens spores collected refrigerated from specimens or of specimens herbarium from obtained spores 2). (Table gans (1989) Ranker follows terminology (1989); Ranker and (1991) (labelled ru ssse oalteohrmjrcae fcelnhis(BS this cheilanthoids of that clades 1.0). major PP confirm other 86%, we the all and to (2009), sister is al. group et bommeriid Windham the sensu of species clade known all include to analysis first the hrce usd h cp fti td u osdrda important an considered but study brandegeei this of of scope synapomorphy the a outside morphology, Perispore character microscope. Meiji a compound on contrast mounted phase micrometer MT5310L ocular an using spores measuring then and ipe hsmasta lhuhtelae of leaves the although that means This simple. ro 18) formerly (1982), Tryon the American of South members the the to of of related members species type as closely 1.0) and PP clade 100%, hemionitid (BS supported rel- strongly closely African are not and is American and South 1.0) atives hypothesized PP its 100%, to (BS related clade bommeriid the of Sporophytes ihntebmeidcae h w cesosof accessions two the clade, bommeriid the Within aeohtsof Gametophytes opooyo prpye,Soe,and Spores, Sporophytes, of Morphology Phylogeny , .ehrenbergiana B. hiate rciea .capensis, C. fractifera, Cheilanthes and Bommeria n all and ohleaochracea Notholaena Notholaena r dnia ooeaohradrbsl supported robustly and another one to identical are r raglradflypnae(erybipinnate) (nearly pinnate fully and triangular are .pedata B. aabrandegeei Baja — omrahispida Bommeria Bommeria $ Bommeria Cheilanthes u hlgntcte Fg )i robustly is 1) (Fig. tree phylogenetic Our Bommeria – n all and , “ h hzm clsof scales rhizome The 0,PP 80%, .elegans B. hlate brandegeei Chelianthes aabrandegeei Baja , species, aabrandegeei Baja Bommeria aabrandegeei Baja .pedata B. ntefed(ocesi (vouchers field the in ( .trichomanoides N. r o resolved. not are ino aooi ramn) Unsterilized Treatment). Taxonomic of tion pce sn aafo ro n Lugardon and Tryon from data using species yRne 18) a oprdamong compared was (1989), Ranker by .aurantiaca C. pce r ih-rw ihadarker a with light-brown are species Bommeria aabrandegeei Baja Bommeria $ ( Results .ehrenbergiana B. ,and .micropteris C. EREE L:BJ,ANWGNSO HIATODFRS473 FERNS CHEILANTHOID OF GENUS NEW A BAJA, AL.: ET GEORGE and , ssrnl eoee samember a as recovered strongly is .9 cosms rnhs hsis This branches. most across 0.99) aabrandegeei Baja Bommeria , r etgnladtechnically and pentagonal are eeoiggmtpye eemade were gametophytes developing .elegans B. , .aureolina N. h eult fnwygerminating newly of sexuality the ae eesae ihPrfl and Parafilm with sealed were lates hiate fractifera Cheilanthes .subpaleacea B. steecpinwt con- with exception the is ’ . .ehrenbergiana B. eim(H7 el 93 in 1963) Hevly 7; (pH medium s u r locmo in common also are but , Tbe2.Telae of leaves The 2). (Table pce ssmaie in summarized is species , .aurea, C. B 7,P .;Fg 1). Fig. 1.0; PP 87%, (BS L)Ds. n hsare thus and Desv.) (L.) and s ntr,sse othis to sister turn, in is, r itntfo that from distinct are group Gametophytes w) h he other three The Sw.). dctdi Additional in ndicated aabrandegeei Baja eaegametophytes, female and , .deltoidea C. and eeclue from cultured were ” and 3 fTynand Tryon of .subpaleacea B. , efblade Leaf . .jaliscana, N. magnification omraele- Bommeria .palmeri C. Bommeria which , — Our Baja Baja Baja are B ro n uadn1991). Lugardon and Tryon of that like The fusion. strand of of spores degrees varying with strands interwoven of finely network a of composed perispores reticulate-cristate and xiie 4salsoe e prnim(i.1.Teothers The ( 1). small (Fig. 32 sporangium produced per spores small 64 exhibited i itiue ln h en oocsoal being occasionally pseudoindu- 2E). undifferentiated (Fig. at to an sium recurved form slightly veins to are intervals that irregular the margins leaf along and submarginal, distributed the 2C). (Fig. half gia midrib least the to at ehrenbergiana margin meria for leaf un- veins the from the the distance with along most contrasts abaxially of This distributed 2A). sporangia (Fig. protected re- margin the leaf by flexed formed lobed a pseudoindusium by covered well-differentiated fully and are and tips vein the near marginally i.1.O h rvosyhpteie eaie of relatives hypothesized previously the brandegeei Of 1). Fig. of species Bommeria five the of four did ( small relatively 64 duced odt hlu Fg ) h aea oiino h multicellular the for in of position meristem lateral 21 The symmetrical 3). a (Fig. day thallus forming cordate ferns), hemionitid in at most in position (as apical tophytes (beginning an plate in initiated prothallial the brandegeei B. the near of mul- cells a marginal base from and initiated plate, was prothallial meristem spathulate ticellular a form to for d proceeded (5 sowing of brandegeei d 18 Baja within germinating began species all of brandegeei among similar qualitatively were lsl eebetoeof and those resemble closely in wide rsn Fg B ,F.Tog prei oeseis all species, some in biseriate ranging in from surfaces, width leaf sparse in abaxial Though their on F). scales have D, bommeriids 2B, always (Fig. are they depending but present re- examined, varies surface is hairs blade and the hair species of the the on density when and attached length The remains the moved. often in embedded that partially surface cells they of leaf unicellular, cluster is bulbous hairs a these from of arise of body true the also Although is 2). this (Table and surfaces, blade leaf their ntal namrel smercltals Continued thallus. asymmetrical markedly a in initially uln ftela Tbe2). pentagonal (Table leaf the the to of contributing outline pinnules, enlarged adjacent are to lobes only pinnae relative proximal are the of pinnae In pinnules pinnules. proximal basiscopic adjacent the than larger of slightly the In pinnules pinnate. connecting thus basiscopic fully and tissue never basal pinnae are leaf proximal blades the of leaf to the leaf wing the small of segment a terminal always is bipinnatifid), to there (pinnatifid lobed elaborately often are species Gametophytes have clade bommeriid expanded newly the of species All Spores h prni of sporangia The l five All .jaliscana N. .pedata. B. .hispida. B. aabrandegeei Baja — nyto( two only , was Bommeria u aidsihl ndvlpetltmn.Spores timing. developmental in slightly varied but l pcmn of specimens All omraehrenbergiana Bommeria .I otat einn tdy3,mrseswere meristems 32, day at beginning contrast, In ). aabrandegeei Baja .pedata B. .I l omris usqetcl division cell subsequent bommeriids, all In ). ,o 4lresoe ( spores large 64 or ), — h ierla clsof scales leaf linear The al aeoht eeomna events developmental gametophyte Early pce xii itntv cclrhison hairs acicular distinctive exhibit species .fractifera C. suuuli hsrgr,wt sporan- with regard, this in unusual is hc ile 2lresoe Tbe2; (Table spores large 32 yielded which , aabrandegeei Baja hiate capensis Cheilanthes ipa ereo tadfso most fusion strand of degree a display , omraelegans Bommeria omraehrenbergiana Bommeria n all and 55 Bommeria aabrandegeei Baja ,3 ag ( large 32 ), m )soe e prnim as sporangium, per spores m) Bommeria and hiate fractifera Cheilanthes Bommeria Bommeria .ochracea N. .pedata B. h xeto within exception The . r itiue sub- distributed are aabrandegeei Baja Bommeria ohleaaureolina Notholaena aabrandegeei Baja oscales to pce htare that species and pce and species pce resulted species xmndpro- examined aabrandegeei Baja sepoo in photos (see ). .deltoidea C. , h basal the , .elegans B. . 0cells 10 game- Bom- most the , Baja Baja ) , Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 MEX 474 ieiodbosrppretgs(BS percentages bootstrap likelihood rwhadcl iiinin division cell and growth e prnimadterrltv ie (s sizes relative their and sporangium per (ARG name species the following code country 3-letter a as shown is phylogeny a with indicated are hlu pt mi imtr ihawl-eeoe ar- well-developed a semicircular symmetrical with nearly diameter, wings. two in and cm mature cushion a 1 chegonial in to resulted up and asymmetry thallus initial this for pensated rhgna rhgnawr is bevda 6dand d 66 at observed first were Archegonia Hevly of archegonia. plates diameter 1 of mm densities 60 (at medium on spores of sowing initial icse ntx r niae nte;caenmsflo ida ta.(2009). al. et Windham follow names clade tree; on indicated are text in discussed Fig. aeohtsof Gametophytes 5 .Pyoeyo hiatodfrsrsligfo aiu ieiodaayi ftecmie plastid combined the of analysis likelihood maximum from resulting ferns cheilanthoid of Phylogeny 1. eio PER Mexico, 5 1 eu USA Peru, in;tikndbace orsodt BS to correspond branches thickened sign); aabrandegeei Baja – 0 prsprpae eeoe only developed plate) per spores 100 5 .brandegeei B. ntdSae,ZAF States, United $ 0 n aeinpseirpoaiiis(PP probabilities posterior Bayesian and 70) 5 ml,L small, htwr rw rman from grown were that aeohtscom- gametophytes 5 ag,c.Dsuso)aeidctdisd prnimsec o ahtxn ao hiatodclades cheilanthoid Major taxon. each for sketch sporangium a inside indicated are Discussion) cf. large, 5 ot fia.Tp pce of species Type Africa). South YTMTCBTN Vlm 44 [Volume BOTANY SYSTEMATIC $ 0 n PP and 80% ’ s 5 retn,BOL Argentina, nifrnitda eulmaturity. sexual male at undifferentiated exclusively male of ( were thalli small The were sowing). gametophytes spores after d 36 form these to began (antheridia from developed that addi- When of mature gametophytes with period. archegoniate plates onto study sown subsequently 12-mo were ga- spores entire tional these the on develop for the not metophytes of did course Antheridia the period. over observation produced continuously be to appeared $ acpiotrsludens Calciphilopteris $ .9 r rvddaogtebace B/P auseult 0 r1.0 or 100 to equal values (BS/PP; branches the along provided are 0.99) .9 egahclpoeac fec oce sdt osrc the construct to used voucher each of provenance Geographical 0.99. Cheilanthes 5 sidctdb natrs ( asterisk an by indicated is oii,CHN Bolivia, ntson steoutgroup. the is shown) (not 5 , atpA hn,CRI China, mi imtr n relatively and diameter) in mm 2 .brandegeei B. , rbcL ,and .micropteris C. 5 ot ia JAM Rica, Costa trnG-R h gametophytes the , .Nme fspores of Number ). aae.Maximum dataset. 5 Jamaica, Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 2019] rmMce n mt 20)ecp for except (2004) Smith and Mickel from uadn(1991). Lugardon ohleatrichomanoides Tryon Notholaena and Tryon of members “ three other The clade. of species type African/Namibian species, brandegeei similar morphologically fractifera lanthes most The morphological grounds. on associated previously been has it which with brandegeei as known formerly species aafo rnilpaigi ntebmeidclade. bommeriid the in it placing unpublished Cranfill of from (2004) data Smith and an Mickel validates by This report species). earlier recognized five all by here resented of brandegeei inclusion the (2009). supports al. strongly well et evidence molecular Windham The summary is by published the ferns 1) with cheilanthoid topology of (Fig. tree overall in tree agrees phylogenetic and supported molecular Our datasets. brandegeei Baja .brandegeei C. Table nti td,w xlrdterltosiso h enigmatic the of relationships the explored we study, this In u oeua hlgntcte Fg )cnim that confirms 1) (Fig. tree phylogenetic molecular Our .Mrhlgclcmaiosars omri en,including ferns, bommeriid across comparisons Morphological 2. sntcoeyrltdt n ftecelnhi ferns cheilanthoid the of any to related closely not is ntebmeidcaea itrto sister as clade bommeriid the in TynadTyn18) oehrwt h South the with together 1982), Tryon and (Tryon group sn ohmlclradmorphological and molecular both using ) uaieSuhAeia eaieof relative American South putative a , Cheilanthes .deltoidea C. ” omasrnl upre ld with clade supported strongly a form Discussion h yeseisof species type the , hiate brandegeei Cheilanthes EREE L:BJ,ANWGNSO HIATODFRS475 FERNS CHEILANTHOID OF GENUS NEW A BAJA, AL.: ET GEORGE ( .micropteris C. and Baja .capensis C. and ntehemionitid the in ) omrahispida Bommeria lse ihthe with cluster , Notholaena Bommeria hri called (herein ’ (1982) s . rw ySsnFwet eipr opooydt rmRne 18) ro and Tryon (1989), Ranker from data morphology Perispore Fawcett. Susan by drawn Chei- (rep- rbcL Baja Baja Baja aabrandegeei Baja n eta mrcnhtpto hiatoddiversity cheilanthoid of 1973). hotspot Tryon and (Tryon American Mexican the Central to confined and are species bommeriid other all given that biogeographically, sense more makes also clade meriid eioadCnrlAeia h rpodapomict pedata triploid The America. Central and Mexico Bommeria hispida Bommeria degeei Although habitat. Sur. by also California and P mountain geographically the Baja a on of Giganta, co-occur side species la these eastern de the Sierra on the range in Peninsula Baja the ain bv 0 Hulr17;Rne n Haufler degeei and Ranker 1979; ele- (Haufler from m known 1990). only 900 are above and vations Mexico mainland in ranges subpaleacea B. Rica; Costa and Nicaragua, Honduras, Guatemala, h atadws oss u ti agl betfo La from absent largely is it but coasts, where west Giganta and east the Vizca and Central the hlgntcpaeetof placement Phylogenetic h rmr igotccaatro the of character diagnostic primary The spiaiyfuda lvtosblw10 ,whereas m, 1000 below elevations at found primarily is group sdsrbtdfo etr n eta eiointo Mexico central and western from distributed is aaaersrce omnaehbtt nmainland in habitats montane to restricted are taxa n l ieseisof species five all and ” ,and srcgie yTynadTyn(92 was (1982) Tryon and Tryon by recognized as omrahispida Bommeria sol on bv 00m l other All m. 1000 above found only is .ehrenbergiana B. ´ ı odsr ein n ok sad off islands rocky and regions desert no omrahispida Bommeria Bommeria nnua hyaesegregated are they eninsula, aabrandegeei Baja cus(i.4). (Fig. occurs aabrandegeei Baja l efilsrtosmodified illustrations leaf All . aemr restricted more have culyocr on occurs actually “ hiate bran- Cheilanthes ntebom- the in sfudin found is aabran- Baja .elegans B. Bommeria , Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 476 nifrnitdpedidsu o ocaigtesoaga .Hisof Hairs F. sporangia. of the Hairs concealing D. not length. pseudoindusium their undifferentiated of (ad hairs much blade leaf for Acicular veins F. D, B, of blade; Hairs leaf B. of sporangia. portion of view Abaxial E. C, brandegeei A, genus. that in found variation Fig. .Cmaio fla agn,soaga itiuin n efbaeidmn in indument blade leaf and distribution, sporangial margins, leaf of Comparison 2. hwn togyrcre,del oe agn omn eldfeetae suonuimlreycneln submarginal concealing largely pseudoindusium well-differentiated forming margins lobed deeply recurved, strongly showing aabrandegeei Baja .C. omraelegans Bommeria omraelegans Bommeria hwn lt nifrnitdmris(opedidsu)adsoagapsto olwn the following position sporangia and pseudoindusium) (no margins undifferentiated flat, showing .E. YTMTCBTN Vlm 44 [Volume BOTANY SYSTEMATIC omraehrenbergiana Bommeria omrahispida Bommeria hwn lgtyadirglryrcre agn omn an forming margins recurved irregularly and slightly showing aabrandegeei Baja . n selected and 5 dxa ar,ab hairs, adaxial Bommeria pce htecmasthe encompass that species 5 bxa ar) A. hairs). abaxial Baja Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 2019] brandegeei ln efmrisadsoagadsrbtdaogteveins the along distributed in with occur sporangia Plants and that margins homoplastic. leaf 1) and plane (Fig. correlated tree are phylogenetic characters molecular these our by suggested with is allied been previously had which group), as supported that to well is discovery group) sister latter the the out- is of representative surprising study typical most this the of of comes One on sporangia. protecting submarginal sporangia margins depending leaf with the recurved margins or families veins, leaf the along plane extending separate have species the into whether ferns much segregated have cheilanthoid so 1970) Pichi-Sermolli correlated, (e.g. strongly authors some be that so to tend or characters pseudoindusium These prominent not. a into differentiated latter recurved is the The whether margin on recurved). based vs. subdivided further (plane is character margins and leaf submarginal) of vs. veins modification the along (extending distribution re- close of indicative ferns. necessarily xeric-adapted among be lationships not may our petioles and erous drought, Tryon of support times in data water important an conserving is to abscission in adaptation Leaf 2). species (Table additional clade three bommeriid in the petioles fractiferous of currence non-fractiferous to and related sam- closely most restricted be to our found Within species the 1). fractiferous the (Fig. of pling, tree members phylogenetic former thoid wherein undeniable is results, “ character petiole our this the of considering near nature homoplastic spaced The evenly base. zones well-defined abscission multiple, transverse by characterized petioles, fractiferous tteopst n ftesetu,rcre margins recurved spectrum, as Notholaena related gen- the distantly in as found of era are pseudoindusia end well-differentiated opposite forming the At andadls aytmsdrn h ogevolutionary long been the have during ferns. features times cheilanthoid of many these history lost clear and is gained It evolution. in margin convergent leaf undifferentiated an ehrenbergiana B. of recurving irregular by the protected sporangia submarginal of appearance occasional iopteris .brandegeei C. hiatodfrsehbtbodvraini sporangial in variation broad exhibit ferns Cheilanthoid Fig. .micropteris C. .Cmaio fgmtpyemrse oiin nA. in positions meristem gametophyte of Comparison 3. mrotrdcae,adin and clade), (myriopterid Bommeria ltrl n B. and (lateral) Bommeria ntoandclade), (notholaenid group Fg E sa vnmr rxmt xml of example proximate more even an is 2E) (Fig. swl si h itnl related distantly the in as well as , epciey(i.1.W lont h oc- the note also We 1). (Fig. respectively , ’ atpclrpeettv fteformer the of representative typical (a ” 16)erirrsrain htfractif- that reservations earlier (1960) s hiate fractifera Cheilanthes r ctee costeetr cheilan- entire the across scattered are Cheilanthes EREE L:BJ,ANWGNSO HIATODFRS477 FERNS CHEILANTHOID OF GENUS NEW A BAJA, AL.: ET GEORGE .deltoidea C. Pellaea Baja .s hmoii clade), (hemionitid s. s. aia) cl bars Scale (apical). bmeidcae.The clade). (bommeriid plai clade), (pellaeid and aabrandegeei Baja .fractifera C. Hemionitis Hemionitis .capensis C. 5 mm. 1 were Myr- .It L. Baja (a . rlne nomr rls ue strands. fused less or more into prolonged prsa opsdo rmwr of framework a of composed as spores uadn(91,wofrtdcmne h pr morphology of spore the documented first who (1991), Lugardon into species this transfer The to in matured. reticulate spores from cristate sequence as developmental coalesce this of to observation continued cristate strands to several the reticulate from as sequence that developmental a species. noted through among features further surface They spore in the producing variation fusion net- strand observed of interwoven degree the finely with a strands of all Gastony of work that composed and perispores observed had Haufler They species (1989). by Ranker thoroughly and (1978b) 1989). studied Ranker were 1991; in genera development Lugardon and and morphology Tryon Spore 1973; Tryon Tryon (e.g. and ferns cheilanthoid of relationships the derstanding ( hairs. cellular one and glands, producing ( deltoidea C. erltdto to related thought formerly species be the Of silicified. heavily and shape rs ida,bttelte ifri en oia in conical being in differ latter the but Windham, & Grusz ubradsz otoeo h orsxa ili pce of species diploid sexual four the of Bommeria those to size and number hiatodfrs n tmyb xlsv oti group. this to in exclusive occur be hairs may other similar it in Somewhat observed and or the ferns, reported of been cheilanthoid members gametophyte not uniting has of clade type initiation bommeriid hair particular lateral The in- 3) meristems. finely and of composed strands, 2), layer Fig. terwoven perispore 2; (Table reticulate-cristate surface a leaf the 2) cluster in bulbous embedded a partially leaves from cells of arising 1) hairs synapomorphies: acicular unicellular, morphological with three share here, sntcoeyrltdto related closely not of is range the per across spores brandegeei scattered Baja small collections 64 19 of from observation sporangium Our for 2018). Scataglini synapomorphy and diagnostic of a species Cheilanthes type provide the may to related sporangium closely taxa in Cheilanthes and 2008) others, al. the et to of species compared two only rare in reported quite been having is just state division character cell this cheilanthoid mitotic ferns Among data). a sexual unpubl. (Windham of in meiosis involves elimination to sporangium prior the per state to spores character due small diploids 32 apomorphic of Another production the 2016). 32 al. yield Haufler unreduced; taxa et are spores apomictic the whereas because larger spores 1986), even (usually larger al. 64 et Barrington produce (see generally al. the polyploids et with (Sigel Sexual clade sporangium, particular 2011). the per on spores dependent size sexual small spore actual 64 for is state species character diploid plesiomorphic the cheilanthoids), including leptosporangiates of clade (the ferns polypod Among relative relationships. cheilanthoid their into reminds insights and critical provide 2015) sporangium can al. sizes per et spores of Schuettpelz number 2012; the al. that et us Li 2011; al. et Sigel ohleaaroia .jlsaa n .ochracea N. and jaliscana, N. aureolina, Notholaena eipr opooyhsbe fgetvlei un- in value great of been has morphology Perispore ebr ftebmeidcae snwycircumscribed newly as clade, bommeriid the of Members eiso eetppr Guze l 09 eke l 2010; al. et Beck 2009; al. et (Grusz papers recent of series A aabrandegeei Baja Fg ;GsoyadHulr1976). Haufler and Gastony 1; (Fig. Fg ) nfc,tepouto f3 ml prsper spores small 32 of production the fact, In 1). (Fig. .s L ta.21;GuzadWnhm21;Ponce 2013; Windham and Grusz 2012; al. et (Li s. s. aen nueto h ld ufcs three surfaces, blade the on indument no have ) eintselegans Hemionitis niae ti eulyrpouigseisand species reproducing sexually a is it indicates aabrandegeei Baja ieiedsrbdisreticulate-cristate its described likewise , Cheilanthes Bommeria w ( two , hiate fractifera Cheilanthes a ioa ntedcso to decision the in pivotal was .s h prsaesmlrin similar are spores The s. s. yipei scabra Myriopteris Rne 99.Tynand Tryon 1989). (Ranker Bommeria hiate capensis Cheilanthes Bommeria Notholaena “ ” olsetrodlets coalescent pce went species aefarina- have ) a multi- has ) n allied and Bommeria (Rothfels (C.Chr.) and Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 478 ld Tbe2.Gmtpyedvlpeti omrisi of is bommeriids in development Gametophyte bommeriid 2). the (Table of more clade members the all to in occurs opposed position, as apical position, typical lateral a in meristem cellular aaae ebru pcmn n ersnsabodsmln falacsil olcin.Bs a iheoein sdwt emsinfo Reb from permission with used ecoregions with map Base a collections. examined accessible specimens all on of based sampling is (2018). broad information a Distribution represents herbaria. 14 and from specimens collections herbarium independent databased 41 of labels herbarium on information location ntedvlpn aeoht,teiiito famulti- a of initiation the gametophyte, developing the In Fig. .Dsrbto of Distribution 4. aabrandegeei Baja nteBj aionaPnnuao eio aapit ersn oriae ae ietyo erfrne from georeferenced or directly taken coordinates represent points Data Mexico. of Peninsula California Baja the in YTMTCBTN Vlm 44 [Volume BOTANY SYSTEMATIC pclcl n ntaino utclua eitmfrom Meristem meristem plate. prothallial multicellular the of a base the of an near of initiation cells marginal absence and the by cell characterized apical is which (1971), Kaur and the “ Ceratopteris -type ” ntecasfcto ytmo Nayar of system classification the in elas well s man Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 2019] aabrandegeei cells Baja of of cluster initiation lateral meristems. bulbous and gametophyte surface, a leaf the in from embedded partially arising trichomes unicellular, of acicular consisting in indument 32 blade (vs. leaf sporangium a per species), spores sexual 64 having in stricto sensu Sw. from Differing sori. con- submarginal strictly completely the deeply cealing pseudoindusium and dissection, well-differentiated leaf pinnate lobed, pen- fully (vs. blades, triangular leaf tagonal) scales, rhizome red-brown concolorous, tde fseisfrom Recent species progresses. of development studies gametophyte young as the persist may in or not may that asymmetry species, bommeriid marked all in thallus a gametophyte in results development 00 ea n are Gal´ y Gal´ Gabriel y and (Gabriel Seral genera 2010; cheilanthoid these in Myriopteris ugs h prto fa nhrdoe ytmlk that like system antheridiogen an in of found operation the prothalli suggest archegoniate mature male of presence of arche- the in development of gametophytes subsequent development and Initial gametophytes (1978a). goniate of Gastony gametophytes and Haufler of development Sexual brandegeei lobes. the development asymmetrical of the of because position apical an in nivea Argyrochosma atod c.50seis n ol aet ecalled be to have would and species) ( Hemionitis 500 latter (ca. the lanthoids of and species 1) type Fig. the including nus in accommodated be longer sasse rmtn ucosn nntrlpopulations. natural in outcrossing promoting system a as a that reported (2011) Baja taxon. dispar- distinctive genus this ecological monospecific the and describe we morphological ity, this acknowledge elevations To at of habitats montane species lower five in occur the all to that contrast in habitats, is desert ( It elevation hot, low 2). occupying (Table by pres- distinguished pseudoindusium the further well-differentiated and a position, of sporangial ence dissection, color, and scale shape rhizome by leaf separable readily is it most observed), the being easily hairs acicular unicellular of indument (the bommeriid clade the of members other with synapomorphies logical Although taxon. this transfer meria 1) remain there options: 1), un- (Fig. viable tree our two of topology and (Schuettpelz the Given a evolution 2018). al. toward cheilanthoid et morphologically progress of of understanding undefinable centuries better two both subverting informative, be would otcoeyrltdto related closely Most ae ntedt rsne herein, presented data the on Based ate brandegeei lanthes brandegeei fri u:MgaeaBy adln sad 1Jan 21 Island, Magdalena Bay, Magdalena Sur: ifornia o.Cu 7 1,p.14 80 T 1890. 104. pl. 215, 17: Club Bot. nov., ida ..ere e.nv T nov. gen. L.O.George, & Windham r2 rc e oopcfcgnst accommodate to genus monospecific new a erect 2) or Bommeria not , lsl olw h atrsin patterns the follows closely hiate brandegeei Cheilanthes aentietfe aea eitminitiation meristem lateral identified not have aabrandegeei Baja DCEtn ida ..ere( L.O.George & Windham (D.C.Eaton) Cheilanthes DCEtn ida ..ere comb. L.O.George, & Windham (D.C.Eaton) aooi Treatment Taxonomic n ealdb afe n eln (1994) Welling and Haufler by detailed and Pi. ida,btta tso appeared soon it that but Windham, (Poir.) aabrandegeei Baja D.C.Eaton). “ sub-lateral Bommeria Cheilanthes uhabodtxnmcconstruct taxonomic broad a Such . EREE L:BJ,ANWGNSO HIATODFRS479 FERNS CHEILANTHOID OF GENUS NEW A BAJA, AL.: ET GEORGE ol nops l oechei- core all encompass would Cheilanthes hiate brandegeei Cheilanthes n21) are Gal´ y Gabriel 2016). an ” .or.btdfeigi its in differing but E.Fourn. ..ao,Bl.Torrey Bull. D.C.Eaton, eitmwsfre in formed was meristem , hrssvrlmorpho- several shares Doryopteris YPE aabrandegeei Baja Bommeria oohltcge- monophyletic A . Baja :M oaccommodate to YPE EXICO nadPrada and an .micropteris C. S , bevdby observed PECIES , Cheilanthes aaCal- Baja . Gaga . Bommeria 00m), 1000 to 5 00m. 1000 a no can and , : Chei- Bom- Baja Baja an ; itiuino h eu nteBj aionaPnnuaof Peninsula California Baja the on Mexico. genus the of distribution n soitdwt ok n hddmcoaiasfrom microhabitats shaded and 30 rocky with associated and ne el urai h uf hsseisi concentrated is of species Vizca as This arid Islands Gulf. well the the Pacific in as in Guarda the Benito, la San de on Angel and and Margarita, 24th Magdalena, peninsula and Cedros, the 30th on the between parallels distributed Mexico; Peninsula, M ig ihrfldegs aegmtpye smaller, gametophytes male laterally. developing Illustrations well- edges; meristem multicellular with with ruffled or maturity ameristic symmetrical at with nearly width) two wings cm and cushion 1 archegonial to developed (up developing large meristem laterally, multicellular with gametophytes male tips. strands. of Gametophytes network vein interwoven finely a near of composed perispore clustered sporangia Spores the recurved the margins, by covered completely segment Sori lobes. lobed, leaflet scales. large prominently of sinuses narrow a at discontinuous occasional forming pseudoindusium well-differentiated recurved, and margins hairs Segment acicular of submarginal abaxial hairs; indument prominent acicular sparse in of than indument adaxial ending hydathodes; larger veins slightly tapering the basiscopic non- only venation anastomosing, basal rounded; base pinnae segments ultimate segment; proximal pinnules; at adjacent distal the nearly pinnate-bipinnatifid ultimate of triangular, pinnules in long, blade pinnatifid as leaf to wide segments; with continuous blade as narrowly wings and colored proximally, lighter costae grooved the distally, adaxially winged rachises and flat- surfaces; distally adaxial on castaneous bundle, grooved vascular longitudinally V-shaped to tened single and a axis) with to perpendicular terete fractiferous grooves abscission proximally multiple than castaneous, margins. (with broader petioles entire slightly scales; bases with rhizome petiole on brown), concolorous scales reddish lanceolate, closely-spaced, to scales rhizome (orange scaly; solenostelic, sdfrsoemaueet;superscript measurements; spore for used aeoht culture). gametophyte acoSnaCtrn nra oPnaCna,2 e 1991, Feb 28 Canoas, Punta to 5462 1991, road Feb on 27 Catarina Santa Canoas, Rancho Punta to road 5426 on Ross Catarina Santa Rancho of 1897, ´ – Plants – Etymology itiuinadEcology and Distribution diinlSeiesExamined Specimens Additional ) ao 19:p.104). pl. (1890: Eaton O); 7 m. 975 h ujc fteseisilsrto ulse ihthe with published illustration species protologue. the of chosen clearly was subject and have rhizome the a to one we attached only leaf the complete and de- is a with specimen collection This Herbarium, lectotype. Eaton the Yale as C. YU.014360 the D. in the All in posited 1889. represented of April are and three March, January, in collected mens GH!, isolectotypes: brandegeei 1890), en- (Eaton fragments; publication one original leaf the of In three RSA!). consisting and sheet, plant of tire side right (YU.014360)!, 1889, nhn 308 Anthony RA;Cdo sad p 1897, Apr 2 Island, Cedros (RSA); 4prsoagu,tiee akabr ihreticulate with amber, dark trilete, sporangium, per 64 RA C;catltrae Wo ut aos 65m of S mi 16.5 Canoas, Punta of NW terraces coastal UC); (RSA, rupestral. rnee s.n. Brandegee –– — i.5 ikladSih(04 7,Fg 75 Fig. 777, (2004: Smith and Mickel 5; Fig. D,M,U,U) oaCetnPit,o ea33m S mi 3.3 mesa on Prieto, Creston Loma US); UC, MO, (DS, a yiidbsdo he rnee speci- Brandegee three on based typified was h eei name generic The rmrl u o xlsvl nsxa:fe- unisexual: exclusively not but primarily ´ ı oadCnrlDsrseoein Fg 4) (Fig. ecoregions Deserts Central and no Mexico. Rhizomes — lcoyedsgae ee YU here: designated (lectotype — B — nei oteBj California Baja the to Endemic AJA Sprcit*idctsspecimens indicates * (Superscript hr-reig horizontal, short-creeping, C Baja ALIFORNIA G rnee 2251 Brandegee niae pcmn sdfor used specimens indicates eest h endemic the to refers ersIln,Mar Island, Cedros : S,U*;San UC*); (SD, Cheilanthes od&Ross & Boyd Leaves od& Boyd Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 480 suonui.Daig ySsnFwetbsdo A: on based Fawcett Susan by Drawings pseudoindusia. Fig. 5. aabrandegeei Baja .Rioesae .Aiua arfo bxa efsrae .Saefo bxa efsrae .Hbt .Dti fsoagaand sporangia of Detail E. Habit. D. surface. leaf abaxial from Scale C. surface. leaf abaxial from hair Acicular B. scale. Rhizome A. . ign n hms176 Thomas and Wiggins YTMTCBTN Vlm 44 [Volume BOTANY SYSTEMATIC S) ,C D: C, B, (SD); emn&Bra 28854 Berian & Rebman S) E: (SD); oa 23952 Moran (SD). Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 2019] nee a,1 p 1947, Apr 16 Bay, Angeles 1949, Apr s.n 1889, Apr 4 Island, Benitos 5 ,1 p 2010, Apr 13 m, 351 L & Salazar nlwrsoe flmsoecif,2.66 14502 l.35m 3Apr 23 m, 365 alt. -114.59022, 29.2696, and lake cliffs, dried 2017, limestone of of side NW slopes on lower canyon on small in Chapala Laguna and Isolote ut rea 7Ot1959, Oct 17 Prieta, 1967, Punta Nov 15 Seca, Chapala Laguna Rancho 5889 al. et Thorne 1982, arntn .S,C .Prs n .A akr 96 ytmtcinferences Systematic 1986. Ranker. A. T. and Paris, A. C. S., D. Barrington, MDW. and LH, T-TK, and KMP, LOG, KMP, Editing: LOG, and Preparation: Review, Draft Writing, Vi- MDW; Original MDW; Writing, Supervision: In- KMP; KMP; MDW; sualization: Administration: and KMP Project Acquisition: LOG; Funding vestigation: T-TK; Analysis: Formal T-TK; and Biology Systematic NSF MDW. an and by KMP to part (DEB-0717398) in award Inventory data. supported georeferenced Biodiversity our was from 4 work Fig. in This map distribution the produced who 2009, Oct 20 m, 26.58489, mesa, 235 of top alt. to -112.72208, road old along Juanico, San and Ignacio San Laguna ta.3980 al. et 1977, Mar 17 3480 Pray m, 1983, 105 Apr alt. 13 m, Refugio, 30 Puerto of SE km 23952 6 peak, of base S IH S* D C S) roo55m S fLsMro,at 5 m, 450 alt. Morros, Los of 1970, ESE Mar mi 5.5 29 arroyo US*); UC, SD, RSA*, MICH, DS, Cerro of W ridge slope, ePaeo,4Mr1992, Mar 4 Placeros, de 62594 Breedlove s.n. 1889, Jan itm el iraVizca Sierra la 1947, de Sistema May 11 Clara, Santa de odt oaio 8Fb1962, Feb 1935, 18 Feb Rosarito, to 19 road 1946, piles, Oct at 30 road m, 1935, 110 of Feb S 23 mi Prieta, 0.25 Punta Prieta, Punta of 7649 Wiggins E mi 9 Angeles, nweg h eeostlnso ua act h rae h line the created who Fawcett Susan of pro- of for talents drawings Haufler generous Christopher the thank knowledge of also spores We maps. viding base of California spores fresh Baja interest acquiring their for purposes. in SD illustration at assistance Hains for Aerne and Layla loans and facilitating Rebman Jon thank for to VT also thanks Special at We staff studies. and DNA their curators our of loans the for for permission YU and sampling US, and UC, SD, collections RSA, MSC, MO, MICH, MEXU, IND, 26.97842, Ignacio, San of W 2014, Abreojos, Oct 7 Punta m, of 105 alt. N -113.48098, Clara, m, 450 Santa alt. 1973, Sierra Feb 9 1973, Azul, Feb Cerro of 7 slope m, 180 alt. Malarrimo, 1973, Arroyo above Feb 3 m, 375 alt. 1971, Nov 3 m, 975 18934 alt. Mechudo, Cerro of ridge N Giganta: la de Sierra aii lp fSer ePaeo,3Jl1985, Jul 3 Placeros, de Sierra of slope Pacific oadcatfo er lno 2.27,-1.10] l.25m Feb 9 Bah m, to 245 1 Highway alt. -114.71603], [29.42777, Blanco, 1962, Cerro from coast toward 1910, ´ ´ S) ersIln,NrhByadhlsdsiln rmlgtos,3 lighthouse, from inland hillsides and Bay North Island, Cedros (SD); . ocpulzto:LG M,adMW aaCrto:L and LH Curation: Data MDW; and KMP, LOG, Conceptualization: etaktehraimcrtr n tf tBY A,D,DK,GH, DUKE, DS, CAS, BRY, at staff and curators herbarium the thank We G,RA;sepPcfcsoecno fSer lcrs u 1986, Jul 3 Placeros, Sierra of canyon slope Pacific steep RSA); (GH, rmsoeadsoaesz nteferns. the in size 149 stomate and spore from idre l 10 al. et Wilder emne l 33108 al. et Rebman ign hms176 Thomas & Wiggins G,MC,S,U,U*;Nsoeo Epa,Pcco eSnaClara, Santa de Picachos peak, SE of slope N US*); UC, SD, MICH, (GH, hee6875 Shreve S*;Cdo sad Ca Island, Cedros (SD*); – rnee s.n. Brandegee 159. RA;Ca (RSA); (SD* asn6119 Dawson opez ´ aabrandegeei Baja oa 17084 Moran D,MC,U) ne n fEcnioCno 11m of S mi 11.1 Canyon Escondido of end inner US); MICH, (DS, G CS IH) ermuho uroNeoCno,Sierra Canyon, Nuevo Puerto of mouth near MICH*); (CAS, ;Esnd,N n fLgn lCprl fLaguna of W Caporal, El Laguna of end NW Ensenada, ); 5262 ´ ı BY S) . iS fLsArsrs ln odto road along Arrastras, Las of SW mi 7.2 RSA); (BRY, adelos MC,M,U) a iSo oaia 6Jn1988, Jan 26 Rosarita, of S mi 8 ca. US); MO, (MICH, omrasubpaleacea Bommeria brae ta.s.n. al. et Oberbauer – ~ n´ aaa ta.5389 al. et Salazar ign 11325 Wiggins 167 nLzr,2.13,-1.34,1 p 1997, Apr 17 -115.03944, 29.41639, Lazaro, on S) ead ert lnoc.Cet a Jos´ San Cuesta ca. Blanco Cerrito de Mesa (SD); oa eel19679 Reveal & Moran G,U) adln sad 1Jn1889, Jan 21 Island, Magdalena US); (GH, ne,at 0 ,2 p 1966, Apr 20 m, 700 alt. Angel, relv 72761 Breedlove uhrContributions Author ´ (SD). RA;Vled gaAag,1 iWo Los of W mi 15 Amarga, Agua de Valle (RSA); ´ ı o e 1947, Dec 3 no, (SD* absns.n. Harbison rnee s.n Brandegee nee,2.21,-1.80,at 1 ,2 Mar 20 m, 615 alt. -114.28306, 29.22917, Angeles, S* C;Isla UC); (SD*, Acknowledgments ign 7716 Wiggins ign 15080 Wiggins ´ ieaueCited Literature htwr sdi i.5 n ioa .Hay M. Nikolai and 5, Fig. in used were that S) a 4k Wb i rmtr f of off turn from air by NW km 24 ca. (SD); — ety7701 Gentry G ign 16738 Wiggins ~ n´ ;R lCieo l.22m 8Arl2010, April 18 m, 212 alt. Chileno, El R. ); B nd aMn,[8314 15203,alt. -115.21023], [28.35164, Mina, la de on EREE L:BJ,ANWGNSO HIATODFRS481 FERNS CHEILANTHOID OF GENUS NEW A BAJA, AL.: ET GEORGE AJA D,RA C;5m fSnBraon Borja San of W mi 5 UC); RSA, (DS, emn&Dlail 18465 Delgadillo & Rebman emn&Bra 28854 Berian & Rebman C (SD* S*;Wo etnHl,4Jl 1979, July 4 Hill, Kenten of W (SD*); D,S) Isla SD); (DS, ALIFORNIA U) ut rea n.d., Prieta, Punta (UC); . CS;3 iS fBah of SE mi 35 (CAS); U) fRnh eqia,alt. Mesquital, Rancho of S (UC); ety7421 Gentry o oprtv td.W ac- We study. comparative for ne el ura lp,W slope, N Guarda, la de Angel G CS S;ra oBah to road US); (CAS, ´ D,RA,U) er Tordillo, Cerro UC); RSA*, (DS, oa eel19978 Reveal & Moran ;9m fPnaPit,1 Feb 19 Prieta, Punta of E mi 9 ); relv 62331 Breedlove aabrandegeei Baja ign 20911 Wiggins RA,S*;Neg fmesa of edge N SD*); (RSA*, D) rooSnJos´ San Arroyo (DS); S mrcnFr Journal Fern American oa 19897 Moran UR adln sad 17 Island, Magdalena : ne el ura N Guarda, la de Angel ´ MC,RSA* (MICH, oa 12906 Moran D) . iSof S mi 2.5 (DS); CS;Picachos (CAS); n furnishing and S) between (SD); S* S;N US); (SD*, (SD). ´ ı Tortugas, a Brattstrom S* US); (SD*, Brandegee ´ ı elos de a ,1 mi 17 e, G Rebman ,UC*); Moran Moran (CAS, ,alt. e, 76: ek .B,M .Wnhm .Ytkeyh n .M re.21.A 2010. Pryer. M. K. and Yatskievych, G. Windham, D. M. B., J. Beck, afe,C .17.Aboytmtcrvso of revision biosystematic A 1979. H. C. Haufler, iehrt .L,J .Rhe,S .Rsel .C eiyr,and Yesilyurt, C. J. Russell, J. Gal´ y S. Gabriel Rohwer, G. J. L., ac- high W. with Eiserhardt, alignment sequence Multiple MUSCLE: 2004. C. R. Edgar, fern. new A 1890. C. D. Eaton, afe,C .adC .Wlig 94 nhrdoe,dr pr ger- spore dark Antheridiogen, 1994. Welling. B. C. and H. C. Haufler, ikl .T n .R mt.20.TePeiohtso Mexico. of Pteridophytes The 2004. Smith. R. phylogenetic A. and for T. J. Testing Mickel, 1996. Kellogg. A. E. and J. R. Mason-Gamer, Link-P´ 2012. Windham. D. M. and Pryer, M. K. F.-W., Li, spore of implications Systematic 1978b. Gastony. J. G. and H. C. Haufler, breeding the and Antheridiogen 1978a. Gastony. J. G. and H. C. Haufler, the Deciphering 2009. Pryer. M. K. and Windham, D. M. L., A. Grusz, monophyletic a Toward 2013. Windham. D. M. and L. A. Grusz, (Pteridaceae: ferns Cheilanthoid 1998. Rollo. R. D. and J. G. Gastony, aso,A 04 lVe:Afs n ihwih lgmn iwrand viewer alignment lightweight and fast A AliView: 2014. A. Larsson, are Gal´ y Gabriel el,R .16.Aattoso hiatodfrst eetenviron- desert to ferns cheilanthoid of Adaptations 1963. H. R. Farrar, Hevly, R. D. Sessa, B. E. Schuettpelz, E. Pryer, M. K. H., C. Haufler, 2019. Windham. D. M. and Huiet, L. Kao, T.-T. Pryer, M. K. O., L. George, analysis sequence 5.0.1 v. Sequencher® 2011. Corporation. Codes Gene atn,G .adC .Hulr 96 hoooenmesadapo- and numbers Chromosome 1976. Haufler. H. C. and J. G. Gastony, ils .M n .J ul 93 neprclts fbosrpiga a as bootstrapping of test empirical An 1993. Bull. J. J. and M. D. Hillis, atn,G .adD .Rlo 95 hlgn n eei circum- generic and Phylogeny 1995. Rollo. R. D. and J. G. Gastony, ikarc,R .B 07 netgtn h oohl of monophyly Investigating the 2007. B. E. Kirkpatrick, R. ilisfrtapoc oseisdlmtto n interpreting and genus fern Botany delimitation the tematic in species evolution to polyploid approach diploids-first rodArboretum Arnold from red ntecneto hlgntcaayi fcelnhi ferns. cheilanthoid of analysis Botany phylogenetic a of context the in of Region. the Floristic in Cape ferns cheilanthoid Greater of radiations for Evidence 2011. Schneider. H. throughput. high and curacy 104. pl. iain n ucosn ehnssin mechanisms outcrossing and mination, h e okBtnclGarden Botanical York New the (Gramineae). Triticeae tribe Biology the in Systematic sets data molecular among conflict geography. Adiantopsis 845 from segregated Botany of Journal American 241 3: in morphology genus fern 1594 the in system the (Pteridaceae). in polyploids apomictic of origins PhytoKeys 10.5061/dryad.qj76320. lanthes adjacent and lines. 131 generic States 17: of reassessment United phylogenetic molecular southwestern A Mexico: the in Cheilanthoideae) dtrfrlredatasets. large for editor 32 pilosa Cheilanthes ln ieccei ih fcneprr research. ments. contemporary of light in 2016. 928 cycle Windham. vascular D. homosporous life M. the and Revising plant Jr, gametophyte: Watkins single the E. and J. Sex Schneller, J. J. Moran, R. from: Data https://www.genecodes.com/ Michigan. Arbor, sequencher. Ann software. h eurcinadrcrusrpinof recircumscription and resurrection The ii ntefr genus fern the in mixis ehdfrassigcniec npyoeei analysis. phylogenetic Biology in confidence assessing for method citoso hiatodfrs(trdca:Celnhiee infer- Cheilanthoideae) (Pteridaceae: ferns cheilanthoid of scriptions rz .A,L .Wto,adR .Hce.21.Rdfnto of Redefinition 2011. Hickey. J. R. and Watson, E. L. A., M. erez, – ryohsanivea Argyrochosma – – 38. 860. – 937. 1601. – 2 504 32: – .l Peiaee.DydDgtlRpstr.https://doi.org/ Repository. Digital Dryad (Pteridaceae). l. s. 256. ora fteAioaAaeyo Science of Academy Arizona the of Journal 2 182 42: 144. n .M n .Paa 00 aeoht fteAda fern Andean the of Gametophyte 2010. Prada. C. and M. J. an, n .M 01 aeoht eeomn n reproduction and development Gametophyte 2011. M. J. an, rbcL 2 49 32: F´ Baja Taxon e(trdca) ytmtc,dvriiain n bio- and diversification, Systematics, (Pteridaceae): ee uloiesequences. nucleotide – mrcnJunlo Botany of Journal American 5 223 35: – 518. 192. Bommeria – e oopcfcgnssgeae from segregated genus monospecific new A : 64. 0 445 60: od.(Pteridaceae). Goldm. 5 524 45: 0 1255 60: Cheilanthes – (Pteridaceae). 234. Bommeria – – Bioinformatics n eae engenera. fern related and 476. ultno h oryBtnclClub Botanical Torrey the of Bulletin 545. – 1 616 81: Bommeria 1268. uli cd Research Acids Nucleic 8 1 88: (Pteridaceae). Taxon . – Biotropica 621. – mrcnFr Journal Fern American 1055. . Biologia 6 1 26: aainJunlo Botany of Journal Canadian hiate yavapensis Cheilanthes 0 3276 30: Astrolepis 6 1636 96: mrcnFr Journal Fern American – 15. Myriopteris Bommeria :1 8: 6 50 66: Bommeria Gaga ytmtcBotany Systematic – :164 2: – 3278. – 11. Pellaea (Pteridaceae). e engenus fern new a , 1645. – ytmtcBotany Systematic 54. 2 1792 32: – (Adiantaceae). . 175. Bioscience (Pteridaceae). ora fthe of Journal (Pteridaceae) 5 341 85: Cheilanthes eor of Memoirs Systematic Systematic complex – 7 215, 17: 1797. – Chei- Aliso Sys- 360. 100: 37: 56: 66: : Copyright (c) American Society for Plant Taxonomists. All rights reserved. Delivered by Ingenta to IP: 152.3.102.254 on: Sat, 28 Nov 2020 16:11:30 P .21.Acmuiydrvdcasfcto o xatlcpye and lycophytes extant for classification community-derived A 2016. I. PPG aa,B .adS ar 91 aeohtso oopru ferns. homosporous of Gametophytes 1971. Kaur. S. and K. B. Nayar, 482 akr .A 99 pr opooyadgnrcdlmtto fNew of delimitation generic and Tracer morphology 2014. Spore Drummond. 1989. J. A. A. T. and Ranker, Xie, D. Suchard, A. M. A., Rambaut, oc,M .adM .Saaln.21.Frhrpors oadthe toward progress Further 2018. Scataglini. A. M. and M. M. Ponce, of names family the of catalogue provisional A 1970. G. E. R. Pichi-Sermolli, between relationships Phytogeographic 2001. Smith. A. and C. R. Moran, cutpl,E,A aia .Pao .Y ia,adG asivc.2014. Yatskievych. G. and Hirai, Y. R. Prado, J. Davila, A. E., Schuettpelz, Pryer. M. K. and Windham, D. M. Huiet, 400 L. from Schneider, H. inferred E., phylogeny Schuettpelz, Fern 2007. Pryer. M. K. and E. Schuettpelz, neotropical of systematics Phylogenetic 1990. A. T. Ranker, cutpl,E,G ohn .M re,C .Rtfl,J Prado, J. Rothfels, J. C. Pryer, M. K. Rouhan, G. E., Schuettpelz, S c h u e t t p e l z ,E . ,K .M .P r y e r ,a n dM .D .W i n d h a m .2 0 1 5 .Au n i f i e da p - ohes .J,M .Wnhm .L rs,G .Gsoy n .M Pryer. M. K. and Gastony, J. G. Grusz, L. A. Windham, Darling, D. M. A. J., C. Ayres, Rothfels, L. D. Mark, der van P. Teslenko, M. F., Ronquist, in combination new 2018. A P. J. 1990. Rebman, Haufler. H. C. and A. T. Ranker, ea,A n .M are Gal´ y Gabriel M. J. and A. Seral, ie,E . .D ida,L ue,G asivc,adK .Pryer. M. K. and Yatskievych, G. Huiet, L. Windham, D. M. M., E. Sigel, ro,R .adA .Tyn 92 en n lidPat,wt Special with Plants, Allied and Ferns 1982. Tryon. F. A. and M. R. Tryon, evolutionary and tropical spores, Geography, in 1973. Tryon. speciation F. A. geographic and M. and R. Tryon, areas America. Endemic South and 1972. Central R. from ferns Tryon, of species New 1960. R. Tryon, 1991. Lugardon. B. and A. Tryon, (*and parsimony using analysis Phylogenetic PAUP* 2003. L. D. Swofford, ferns. 175 oaia Review Botanical 304 ora fBotany of Journal World http://tree.bio.ed.ac.uk/software/tracer/. 1.6. v. hsso ot mrcnspecies. American South on phasis of delimitation pteridophytes. living pteridophytes. African-Madagascan and neotropical oeua hlgntcadmrhlgclafnte of affinities morphological and unsampled phylogenetic previously Molecular of affinities the and genera. relationships Assessing Pteridaceae: family overall fern the of phylogeny molecular A 2007. genes. plastid three and species leptosporangiate 712 vonoids. Bommeria .A ude .D ida,R .Mrn n .R mt.2018. Smith. R. A. genera? and fern Moran, many C. too R. there Windham, Are D. M. Sundue, A. M. raht aooi eiiaini h engenus fern the in delimitation (Pteridaceae). taxonomic to proach senae atrso vltoaycnegnei ei-dpe ferns. xeric-adapted in convergence evolutionary monophyletic of patterns a Toward 2008. model space. and model inference large a phylogenetic across Bayesian choice Efficient 3.2: MrBayes H¨ S. 2018). May accessed (last http://bajaflora.org Museum. History (Adiantaceae). 01 pce eainhp n aiaeouini h cheilanthoid the in evolution farina genus fern and relationships Species 2011. triphylla eeec oToia mrc.NwYr:Springer-Verlag. York: New America. Tropical to Reference Society Linnaean the of Journal Botanical 145 Pp. Ferns of ferns. Classification cheilanthoid the in relations ferns. American Rhodora Springer-Verlag. Associates. Sinauer Sunderland: 4. v. methods), other – – – ha .Lre,L i,M .Scad n .P ulebc.2012. Huelsenbeck. P. J. and Suchard, A. M. Liu, L. Larget, B. ohna, 186. 351. 724. (Pteridaceae). ora fSseaisadEvolution and Systematics of Journal eints Gymnopteris Hemionitis, oeua hlgntc n Evolution and Phylogenetics Molecular Peiaee Polypodiopsida).(Pteridaceae, 2 1 62: ytmtcBotany Systematic Ainaee ae nmrhlg,alzms n fla- and allozymes, morphology, on based (Adiantaceae) Argyrochosma – h lr fBj California Baja of Flora The 10. ytmtcBotany Systematic mrcnFr Journal Fern American Cheilanthes 7 295 37: Biotropica 6 297 76: Taxon d.A .Jry .A rbe n .A Thomas. A. B. and Crabbe, A. J. Jermy, C. A. eds. , Webbia – – 396. (Pteridaceae). 306. n 06 aeohtcpaeof phase Gametophytic 2016. an. 3 258 63: :121 4: 5 442 15: Celnhiee trdca) ihem- with Pteridaceae), (Cheilanthoideae; 5 219 25: ,and prso h Pteridophyta the of Spores Notholaena – 0 629 40: – Taxon 131. raim,Dvriy&Evolution & Diversity Organisms, – 264. Bommeria ytmtcBiology Systematic 453. – 0 1 80: 297. oaiaComplutensis Botanica a ig:SnDeoNatural Diego San Diego: San . ytmtcBotany Systematic 7(up.1). (Suppl. 67 – 7 473 67: 644. 4 563 54: – – 3. 5 in 153 Peiaee:Resolving (Pteridaceae): (Adiantaceae). 4 1172 44: – Taxon 480. – 603. h hlgn and Phylogeny The – 1 539 61: Hemionitis 1185. 6 1037 56: Pentagramma e York: New . Brittonia YTMTCBTN Vlm 44 [Volume BOTANY SYSTEMATIC 6 554 36: Doryopteris 0 63 40: Adiantum Bommeria Taxon American – 542. – 1050. – 564. – and 18: 53: 57: 70. ohleatrichomanoides Notholaena 89-285 Gastony & Yatskievych JQ855909. Nayarit, EU268675. EU268778, KF961864. Arbel´ KF961803, KF961739, #6158, UC, EU268690. ynpei marantae gymnopteris dlesex, Heredia, EU268683. Rica, EF452145, MK020110, #3709, MO, MK020128. Lima, Peru, Tryon, R.M. 2187 MK020126. .brandegeei B. Sur, fia otenCape, Northern Africa, MK020125. Puebla, MK020115, MK020106, Mexico, MK020124. MK020114, Maxon, MK020105, #6940, DUKE, EU268671. Arizona, USA, MK020123. U27729, MK020104, O 33,E286,E426,EU268711. EF452161, EU268763, #3736, MO, Hidalgo, Mexico, MK020121. Underw., MK020113, MK020102, enDAdtbs ubr(ena.ilg.ueeu,Gnakac- for GenBank numbers (fernlab.biology.duke.edu), number cession database DNA Fern analys phylogenetic division, political country, molecular the in Phylogenetic the for Approaches Algorithm Genetic 2006. J. D. Zwickl, afe,Mxc,Oaxaca, Mexico, Haufler, MK020122. U19497, MK020103, 31,E284,E425,EU268688. EF452150, EU268741, #3510, Cultivated Germany, H.Schneid., JQ855913. EF452162, Virginia, JQ855925, from originally - Cultivated USA, ida,M . .Hit .Shetez .L rs,C .Rothfels, J. C. Grusz, L. A. Schuettpelz, E. Huiet, L. D., M. Windham, asivc,G n .L .Arbel´ A. L. A. and G. Yatskievych, eiyr,J . .Barbar´ T. C., J. genus Yesilyurt, fern new The 2010. Schneider. H. and C. J. Yesilyurt, iue1. following Figure 2 and 1 numbers brandegeei 622 Castillo & Torrico Appendix Outgroup: Ingroup: C 12,M000,M001,MK020127. MK020117, MK020108, #1126, UC, , ieiodCiein hD israin utn ea:TeUniver- The Texas. Texas: of Austin, sity dissertation. Maximum Ph.D. the Criterion. Under Datasets Likelihood Sequence Biological Large of Analysis pce opee ncelnhi ferns. cheilanthoid 128 and in plastid demystify Using complexes and 2009. boundaries species generic Pryer. redraw M. to K. sequences and DNA nuclear Yatskievych, G. Beck, B. J. 120 genus fern the in transfers generic .Gby 05 dniyn h eei iiso h cheilanthoid the of limits generic the Identifying genus 2015. Gibby. M. (Pteridaceae). relv 62594 Breedlove e,Mxc,Oaxaca, Mexico, aez, akr860 Ranker – – 132. 124. ohese l 3118A al. et Rothfels 1 ohleaochracea Notholaena DC ao)Wnhm&LOGog,Mxc,Bj California Baja Mexico, L.O.George, & Windham Eaton) (D.C. yipei marsupianthes Myriopteris Doryopteris hiate micropteris Cheilanthes hiate deltoidea Cheilanthes omrapedata Bommeria ryohsanivea Argyrochosma .Vuhr n eBn ceso ubr o aaused taxa for numbers accession GenBank and Vouchers 1. 2 eio aaClfri Sur, California Baja Mexico, , acpiotrsludens Calciphilopteris ohese l 08-184 al. et Rothfels cutpl 467 Schuettpelz Phytotaxa T 45,E286,E280,EU268710. EU268807, EU268762, #4054, UT, , IH 40,M000,M001,MK020120. MK020112, MK020101, #4508, MICH, , L)KHSig hn,Yunnan, China, K.H.Shing, (L.) O 63,H869,H864,HQ846493. HQ846445, HQ846396, #6236, MO, , . atpA ohgae734 Mothogoane ohleajaliscana Notholaena Phytotaxa anes353 Saunders L)Ds.var. Desv. (L.) ,H cnie,S usl,A uhm and Culham, A. Russell, S. Schneider, H. a, olco() olcinnumber, collection collector(s), ida ta.544 al. et Windham , Rojas-Mart N,#55 U678 U677 EU268674. EU268777, EU268728, #4515, IND, , asivc atn 89-258 Gastony & Yatskievych :52 7: S. .or. eio Jalisco, Mexico, E.Fourn., (Sw.) aabrandegeei Baja Ho. as.&Arbel´ & Yatsk. (Hook.) asivc atn 89-203 Gastony & Yatskievych rbcL Pi. ida var. Windham (Poir.) uz,SuhArc,Cp Province, Cape Africa, South Kunze, UE 65,M001,MK020119, MK020111, #6553, DUKE, , omrahispida Bommeria UE 37,E282,EF452142, EU268725, #3174, DUKE, , – w,Agnia Misiones, Argentina, Sw., 2:101 221: – omraelegans Bommeria , hiate capensis Cheilanthes 59. rgnunknown, origin UE 53,E429,KC984525, EF452098, #5137, DUKE, , trnG-R e.20.Anwseisadthree and species new A 2008. aez. 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