Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 94 ag17;Hulre l 95,b,temnpyyo the taxa of diploid its monophyly among the Lang relationships b), and phylogenetic 1995a, and al. Lloyd the complex et 1961a; Haufler of 1971; Shivas members Lang 1950; 1964; polyploid 1947, the (Manton of complex origins the and 1995b). al. et Haufler loerpodtxndrvdfo h ili progenitors diploid Asian the glycyrrhiza and European from North derived the taxon to allotetraploid only applied is s.) (s. stricto aeo ulcto etme ,2014 3, September publication of Date 10.1600/036364414X683921 DOI © Botany Systematic 95;Shao 01.Tdy h name the 1991; Today, 1971; Zhongren al. 2001). et 1969, Haufler and Schmakov 1993; al. Lang 1995b; Haufler et Haufler 1991; 1964; 1991; Windham 1963; and Lang Haufler Smith Lang and and and Whitmore Lloyd Taylor 1964; 1963; Valentine (Lloyd worldwide the constitute complex as that hybrids, species, sterile allohexaploid numerous ten as one well about and of allotetraploid, recognition six the diploid, to led have investigations quent of cytotypes to vulgare European status reproducing P. sexually specific three assigned the they of studies, each confu- these taxonomic the on of Based source sion. one as implicate reticulation to of first history the a were b) (1961a, Shivas and (1950) experiments, Manton hybridization and studies cytotaxonomic splitUsing to demonstrated, began and were detail researchers geography greater with in correlations examined strong were their 1990) and al. 1961a et Shivas Hennipman by (summarized characters cryptic these rhizome varieties. taxonomic As as and treated ornamentation, or dismissed either and were flavor, size spore shape scale coloration, rhizome and type, including indument morphology, shape, leaf in in differences and variations Eurasia Subtle northern of America. much spanning North species single a be to taxon- centuries, 20th and considered 19th omists the of much For reticu- evolution. of late patterns complex that with ferns morphology cryptic of combine examples best-studied the among are podiaceae) oyih 04b h mrcnSceyo ln Taxonomists Plant of Society American the by 2014 Copyright 1 ept rgesi eovn h eiuaerelationships reticulate the resolving in progress Despite ebr fthe of Members eateto ilg,Dk nvriy o 03,Booia cecsBidn,Dra,NrhCrln 70 .S A. S. U. 27708 Carolina North Durham, Building, Sciences Biological 90338, Box University, Duke Biology, of Department u oeaeetmtst esespeiu yohss n opooenwhptee,aottehsoia vnsta hpdtediversity the shaped that events historical use the the we of about Finally, species hypotheses, Pleistocene. diploid new the the propose through of Miocene to late and distribution the hypotheses, geographic from previous current diversifying reassess and species to diploid estimates extant age of node majority our the with complex, the of lineages yoaooi ok yrdzto xeiet,adioyesuiscnutddrn h 0hcnuy ept osdrbeeffort, considerable Despite century. species 20th the single of the a members during considered diploid conducted Once the the studies of among cluster. phylogeny isozyme relationships diploids-only species evolutionary and reticulate the experiments, America, highly however, North hybridization and and work, confusing Eurasia cytotaxonomic a northern of generated much have spanning and morphologically differentiated osldt,w siaea al icn rgnfrthe for origin Miocene early an estimate we data, endemic fossil Asian the to sister ( are loci The plastid clades. four well-supported from four comprising data sequence Using stricto. sensu hlgn,Dvrec ieEtmts n hlgorpyo h ili Species Diploid the of Phylogeography and Estimates, Time Divergence Phylogeny, Abstract— Keywords— Mno 97 90 91 hvs16a ) Subse- b). 1961a, Shivas 1951; 1950, 1947, (Manton ..Etnand Eaton D.C. 21) 94:p.1042–1055 pp. 39(4): (2014), rnM Sigel, M. Erin 2 eateto clg n vltoayBooy nvriyo ass 20SnyieAvenue, Sunnyside 1200 Kansas, of University Biology, Evolutionary and Ecology of Department The iegnedtn,Hwia sad,Mcrnsa lsi eunedt,Pleistocene. data, sequence plastid Macaronesia, Islands, Hawaiian dating, Divergence .vulgare P. oyoimvulgare Polypodium oyoimvulgare Polypodium fthe of .vulgare P. h yeseisof species type the , .sibiricum P. 1,3 .vulgare P. luooipi makinoi Pleurosoriopsis ihe .Windham, D. Michael ope Plpdaee opie elsuidgopo entx hs ebr r cryptically are members whose taxa fern of group well-studied a comprises (Polypodiaceae) complex oyoimvulgare Polypodium 3 nosgeaespecies. segregate into uhrfrcrepnec ([email protected]) correspondence for Author ope n eae pce ots rvoshptee ocrigrltosiswithin relationships concerning hypotheses previous test to species related and complex aot al arne ass605U .A. S. U. 66045 Kansas Lawrence, Hall, Haworth ilv Sia 1961b; (Shivas Sipliv. .vulgare P. .cmlx(Poly- complex L. .vulgare P. omnctn dtr eleGoertzen Leslie Editor: Communicating Polypodium sn iegnetm nlssicroaigpeiul eie g osrit and constraints age derived previously incorporating analyses time divergence Using . ope srsle ssse oteNeotropical the to sister as resolved is complex .vulgare P. .vulgare P. .vulgare P. tA bL matK, rbcL, atpA, .vulgare P. sensu 1 hitpe .Haufler, H. Christopher .vulgare P. L., P. 1042 ope n aeMoeePicn rgnfrtefu ao diploid major four the for origin Miocene-Pliocene late a and complex a ensgeae no1 ili n oyli aaa eutof result a as taxa polyploid and diploid 17 into segregated been has ope (Polypodiaceae) Complex andthethird(cladeC: Ameri North predominantly are eti ssse to sister sug- is studies it molecular gest some relationship endemic; contentious coast American most of North the that Perhaps is diploids a–f). among endemic 1 Hawaiian the (Fig. of dies placement the pellucidum these as P. within well as and clades, among 1943, three Relationships Martens The iv; 1964). midrib. and Wagner leaf iii 1950; 1 the (Fig. sporangia of the surface among tributed adaxial the cambricum on P. hairs In has sporangia. but to homologous the are contrast, that Martens sori glandular ii; often within sterile, and are structures i which 1974), 1 Kott Fig. and Peterson (see 1943; sporangiasters have all the (A) of clade Mediterranean Members the Macaronesia. primarily in Europe, and region, in found is Bobrov) A.E. 95 afe ta.20) hra tesso ta itrto sister as it show others whereas endemic Hawaiian 2000), the al. et Haufler 1995; G: clade toe l 09.Too hs caeA: (clade these 2004; of al. + Two et Windham & 2009). Schneider Haufler 2000; al. 1995b, Ranker et 1995a, and Otto al. Haufler 1980; et Roberts Haufler 1971; Lloyd1995; Lang 1950; 1964; Manton Lang 1; and (Fig. a character by morphological united each shared taxa, diploid provides of clades studies major previous three of for support synthesis A resolved. fully not are oemlclraaye upr itrrltosi between relationship sister comm.; a support pers. analyses molecular Smith, some the R. (A. of species Tejero-Dı recircumscription polyploid and a diploid 20 for need plesiosorum the phylogenetic reveals recent While b). work 1995a, al. et Haufler 1982; Tryon plesiosorum P. lxhsbe osdrdcoeyrltdt h Neotropical the to related closely considered been has plex complex. and nmrhlgclgons the grounds, morphological On .vulgare P. trnG-trnR ´ Pglycyrrhiza zadPceo20;Ln-eae l 02.Whereas 2012). al. et Luna-Vega 2004; Pacheco and ez ru Ot ta.20) tcmrssa estimated an comprises it 2009), al. et (Otto group .glycyrrhiza P. ld C a rnhn,hi-ieprpye dis- paraphyses hair-like branching, has (C) clade al. eentwl eovdb rvosstu- previous by resolved well not were Kaulf., ope eanpol eovd eew ne a infer we Here resolved. poorly remain complex ,w eoee monophyletic a recovered we ), uz ru Crsesn12;Tynand Tryon 1928; (Christensen group Kunze .glycyrrhiza P. 2 + n ahenM Pryer M. Kathleen and .pellucidum P. .californicum P. .amorphum P. .plesiosorum P. .cambricum P. ld G sdvi fsporangiasters, of devoid is (G) clade .scouleri P. Fg ,e afe n Ranker and Haufler e; d, 1 (Fig. a u xedt atr Asia, eastern to extend but can Fg ;Ot ta.2009). al. et Otto f; 1 (Fig. ok rv,awestern a Grev., & Hook. ru n hs,i turn, in these, and group oyoimvulgare Polypodium us.+ Suksd. al.+ Kaulf. .+ L. .vulgare P. .macaronesicum P. .appalachianum P. .appalachianum P. .sibiricum P. .fauriei P. Polypodium 1 complex Christ) com- and P. Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. aahssfudin found paraphyses nti td.Istiae r oiidfo atn 14)a olw:i prniseswt lnua rcoe on in found trichomes glandular with sporangiasters i. follows: as (1943) Martens from modified are images appalachianum Inset P. study. this in of analysis inference from derived trees parsimonious most ( C and etito ie n lsi eunedt yHulre l 19a )adHulradRne 19) ml etudrec ao aeidctsthe indicates name taxon each under text Small (1995). Ranker ( and A Haufler as marked and are b) clades (1995a, recovered commonly al. Three et species. Haufler that for by distribution data geographic sequence generalized plastid and site, restriction Neotropical akr19;Hulre l 00 toe l 09.Tu,even Thus, 2009). al. et Otto 2000; al. et Haufler the 1995; within Ranker the nested that be suggest others may 2009), al. et Otto the 1043 PHYLOGENY POLYPODIUM AL.: ET SIGEL 2014] ³ ae nboegahcpten n opooia yaoopis .Hulre l 19b,drvdfo eei nlsso szm aa .Hauf c. data; isozyme of analysis genetic from derived (1995b), al. with et nodes Haufler (1995a), b. synapomorphies; al. morphological et and patterns biogeographic on based 0 otta upr naprioyaayi of analysis parsimony a in support bootstrap 70% Fig. .vulgare P. .cambricum P. .Cnesstplg fpeiul ulse eainhp o ili pce fthe of species diploid for relationships published previously of topology Consensus 1. oyoimplesiosorum Polypodium and i prniseswtotgadlrtihmsfudin found trichomes glandular without sporangiasters ii. ; ld) rydse ie n oecs etr eittplgcldfeecsrcvrdb pcfcsuis .Hulre l (1995b), al. et Haufler a. studies: specific by recovered differences topological depict letters lowercase and lines dashed Grey clade). .cambricum P. rLF rbcL, trnL-F, .plesiosorum P. ³ 0 otta upr naprioyaayi fclrpatrsrcinst aa .HulradRne 19) oe with nodes (1995), Ranker and Haufler d. data; site restriction chloroplast of analysis parsimony a in support bootstrap 70% .vulgare P. and oiotllnsnx oistiae ersn 100 represent images inset to next lines Horizontal . ru.Tikbaklnsrpeettecnesstplg nerdfo opooia,ioye chloroplast isozyme, morphological, from inferred topology consensus the represent lines black Thick group. rps4 rus(afe ta.1995a, al. et (Haufler groups trnL-trnF lsi eunedt.Ol afe ta.(00 a opeesmln ftetndpodseisincluded species diploid ten the of sampling complete had (2000) al. et Haufler Only data. sequence plastid ope Hulrand (Haufler complex lsi eunedt;f toe l 20) oe ih=10pseirpoaiiyspoti Bayesian a in support probability posterior 1.0 = with nodes (2009), al. et Otto f. data; sequence plastid .plesiosorum P. rbcL lsi eunedt;e afe ta.(00,rltosisbsdo titcnesso 180 of consensus strict a on based relationships (2000), al. et Haufler e. data; sequence plastid group .sibiricum P. oidrjl ta.21)t netgt eainhp within relationships investigate to 2011) al. et Govindarajulu 1). (Fig. question h oohl fthe of monophyly the eew dp dpod-is”apoc Bc ta.2010; al. et (Beck approach “diploids-first” a adopt we Here m m. i.srlprpye on in found paraphyses soral iii. ; oyoimvulgare Polypodium .appalachianum P. .vulgare P. ope a encle into called been has complex ope n ebr fthe of members and complex ld) ( G clade), .macaronesicum P. .gyyriaclade glycyrrhiza P. .amorphum P. v soral iv. ; and ler ), Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ota eu Mce n mt 04 toe l 09.Aaye were Analyses 2009). al. et genus Otto scaly-leaved 2004; the Smith and with with rooted (Mickel allied genus closely that be to Polypodiaceae to the shown in Pleopeltis was clades that were major taxa cies of outgroup other diversity including Seven 1), a (Appendix 2009). represent al. et to Otto selected 2004; al. et (Schneider h oyoica ShetezadPyr20) oehrwith arecentlydescribed together 2009), Pryer and (Schuettpelz Polypodiaceae for dates the divergence calibrated established use ferns. we iconic addition, these In of phylogeny molecular a infer we complex, the to assigned species diploid ten includes that pling upr;BP:Bysa neec otro rbblt;*Tecmie aae a nlzdudrapriinmdl ihec ou ie t own its given locus each with model, partition a under analyzed was dataset combined The * probability; model. posterior best-fitting for inference Bayesian data BIPP: support; indels coded binary to opod eas falc fctgntcdt o hs taxa. these for data to relative cytogenetic closest the the as including of studies clade molecular lack recent a in position a its on based of 2009). al. makinoi because et Pleurosoriopsis Otto ploidy 2004; Smith to and Mickel the to 1995; of related Representatives Moran be 1982; to Tryon shown and subsequently the but to groups, assigned formerly taxa the the in to taxa belonging diploid recognized ten obnd4 ,5 5 46– 27 .976 0.89 75 92 –* 24.6 951 4,452 42 Combined indels trnG-trnR trnG-trnR rbcL matK atpA 39 [Volume the detailing studies the ongoing within formation allopolyploid for of timing foundation and history the reticulate BOTANY SYSTEMATIC lay species; 4) diploid the and of origin the for context biogeographic and the the of with taxa diploid relationship the the sister of hypothesized monophyly plesiosorum P. the its assess and 1) complex to: are goals Our of divergence the Polypodium 1044 ope.UigDAsqec aafo orpatdloci plastid four from data sequence ( DNA Using complex. aiona olwn h auatrrspooo.Oet orplastid Valencia, four (Qiagen, to Kit One silica-dried, Mini protocol. Plant herbarium, manufacturer’s regions, DNeasy from the the following isolated using California) was material DNA fresh or genomic total pled, 2009). al. et Otto ta.2011; on al. based et 2006; al. were et protocols, Schuettpelz from sequencing ESTRNR46F and ( sequenc- amplification and studies amplification as previous for well used as Primers ing, 1). (Appendix samples the trnG-R rtdsqecsaeaalbei eBn Apni 1). gen- (Appendix newly GenBank 115 in The available are GenBank. sequences sets from erated data 2007; sequences Plastid with 2007). al. supplemented Pryer et were Nagalingum from and TRNR22R and TRNG63R, Schuettpelz from atpA ao Sampling— Taxon N xrcinadPatdSequencing— Plastid and Extraction DNA Table Polypodium .vulgare P. ou o fSqecsIcue ie aibeSts%MsigDt etFtigMdlMa Partitions % Mean Model Best-Fitting Data Missing % Sites Variable Sites Included Sequences of No. Locus , ,wr nieyo atal mlfe n eune o ahof each for sequenced and amplified partially or entirely were ), matK tA aK rbcL, matK, atpA, ublt&Bnln xWldnwbthsytt transferred to yet has but Willdenow ex Bonpland & Humboldt .Saitc o h aaesaaye nti td.Msigdt nld ohucranbss(,N n as() eunedt and data Sequence (-). gaps and N) (?, bases uncertain both include data Missing study. this in analyzed datasets the for Statistics 1. rbcL Platycerium , .s,wt atclrfcso the on focus particular a with s., s. SBLF SBL2F SBL5R n ESRBCL1361R and ESRBCL654R, ESRBCL628F, ESRBCL1F, : .s codn orcn tde Shedre l 2004; al. et (Schneider studies recent to according s. s. rbcL complex. ru;2 ne vltoayrltosisamong relationships evolutionary infer 2) group; .plesiosorum P. atpA ,and aeil n Methods and Materials esmld34 sampled We Polypodium .plesiosorum P. STA1F STA3F STA5R and ESATPA557R, ESATPA535F, ESATPA412F, : Mxm)Fmnwsicue nteanalyses the in included was Fomin (Maxim.) oyoimvulgare Polypodium ev,teotru ao otdsatyrelated distantly most taxon outgroup the Desv., 11392785GR+G9 1–– – – – – – 81 56 81 Mkv 90 79 G + 76 K81uf G + G GTR + TVM 0 G + TrN 1.0 8.5 2.7 31 5.9 230 227 232 31 231 964 1,309 624 1,524 29 29 41 24 34 oyoimsanctae-rosae Polypodium Polypodium n the and .vulgare P. trnG-trnR .dulce P. ru Apni ) hs nld two include These 1). (Appendix group trnG-trnR trnG-trnR .s n the and s. s. .vulgare P. ru eeslce ihu regard without selected were group Polypodium ope;3 rvd temporal provide 3) complex; osl(Kvac fossil negncsae)adsam- and spacer) intergenic trnG-R or or Poir. matK eeuie noasnl aiu ieiod(L nlss LS aiu ieiodbootstrap likelihood maximum MLBS: analysis. (ML) likelihood maximum single a into united were negncsae (henceforth spacer intergenic + ERadrL rmKuo from rGLR and fEDR : o ahidvda sam- individual each For ope n iespecies five and complex .plesiosorum P. RGF TRNG43F1, TRNG1F, : pcmn representing specimens Mxn .Cr,aspe- a Chr., C. (Maxon) .subpetiolatum P. .vulgare P. .plesiosorum P. ˇ k20) odate to 2001), ek .vulgare P. .vulgare P. .vulgare P. complex. complexes (Tryon Hook. xlddtefrttomlingnrtoso ahrna burn-in, as run each tree consensus of PAUP majority-rule A using generations samples. generated 32,000 we of million was pool conservative, final two be a in to first resulting each but runs the generations, four The million excluded 2007). one Drummond around and visual- (Rambaut converged were sample v1.5 traces a Tracer parameter with in sample generations ized The four million generations. other with 10 1,000 run all for every were and heated), taken analyses each three variable) Four for cold, values. = rates (one default (ratepr average chains their The at different left variable. be were = to priors coding allowed and equal, were The = partition rates settings. 1, gamma = the = nst best- for rates The model and dirichlet(1,1,1,1) settings. Mkv = gamma statefreqpr the = mixed, for rates model and fitting dirichlet(1,1,1,1) = statefreqpr fmdl cetdb rae,tebs-itn oesfor models best-fitting range the the cluster MrBayes, were accommodate by to Parameters and computing order accepted 2010). CIPRES In models al. partitions. of the et five the Miller on among 2003; unlinked 3.1.2 Huelsenbeck v and (Ronquist MrBayes in implemented above described approach ML analyses. the single-locus the using for analyzed was data) indel coded h ags l cr te ttsissmaie nTbe1.Freach was For dataset 1). each PAUP for Table using tree compiled consensus in majority-rule pseudoreplicate bootstrap bootstrap summarized 1,000 The on statistics datasets. performed were (tree searches having tree score as dataset, identified -ln maxi- was inde- tree optimal two largest ML 20 for “best” a the in single run resulting The with was trees. each, data likelihood analysis as replicates mum Each ten indel model 2001). with best binary-coded searches Lewis the of pendent 1; with (Table partition data model a sequence Mkv and of above, partition a determined analysis: for tions eune rmmlil pcmn of specimens Kunze, multiple dataset. from four-locus combined Sequences one and were datasets datasets single-locus Five four and analysis. 2.1 compiled: from alignment, excluded the v. were to indels gapcode.py appended corresponding were in the of characters implemented binary method These the as 2008). using in (2000), (Ree individual only each Ochoterena (present for and absent Indels Simmons or data. present missing as as coded were coded were of portions locus Unsequenced with each 2005). aligned Maddison manually and (Maddison were 4.05 Codes locus MacClade (Gene each for 4.8 Sequences Sequencher 2005). using Corporation assembled and edited manually were aho h orsqec oiadoepriinfrthe for partition one and loci sequence combined four single for partition the a one partitions: of into data each datasets five (with single-locus dataset combined four This dataset. the allowing concatenate detected, were to incongruences us significant No 1996). Kellogg and upre ( supported The 2012). al. et ( and Hett. & Hennipman (Mett.) aae Apni 1). (Appendix dataset contigua Prosaptia M)a mlmne nGri20o h IRScmuigcluster for computing sets CIPRES data Single-locus the 2010). on al. 2.0 et and likelihood Garli Miller maximum 2006; in using (Zwickl datasets implemented single-locus as the (ML) of each for ducted kieIfrainCiein(I;Tbe1 nPriinidr(Lanfear PartitionFinder in 1) Table (AIC; Criterion Information Akaike .stemaria P. euneAinetadDt Sets— Data and Alignment Sequence hlgntcAnalyses— Phylogenetic h obnddtstwsas nlzduigBysa neec as inference Bayesian using analyzed also was dataset combined The trnG-R rbcL rmteimmiyoshianum Drymotaenium eeaaye sn h etmdla eemndb the by determined as model best the using analyzed were Bav)Ds.and Desv. (Beauv.) ³ eunepriin eeipeetdwt h s 6, = nst the with implemented were partitions sequence 0 otta upr)icmail lds(Mason-Gamer clades incompatible support) bootstrap 70% trnG-R .Pel eecnaeae ntecmie four-locus combined the in concatenated were Presl. C. trnG-R * matk .4013(wfod20)adcmae o well for compared and 2002) (Swofford 4.0a123 v. igelcsdtstwsdvddit w parti- two into divided was dataset single-locus 36 – – 64 83 eaaepyoeei nlsswr con- were analyses phylogenetic Separate * atto a mlmne ihtent= nst the with implemented was partition ne atto a mlmne ihthe with implemented was partition indel .4013(wfod20) h combined The 2002). (Swofford 4.0a123 v. .superbum P. LSBIPP MLBS .fortunei M. Makino, N eunechromatograms sequence DNA ³ .peisrm .rhodopleuron P. plesiosorum, P. eJnh enpa) and Hennipman), & Jonch. de 0 en%Partitions % Mean 70% TMoe Ching), (T.Moore) Microsorum ik( Link trnG-R tA matK, atpA, Platycerium .varians M. atpA trnG-R binary- , ³ rbcL 0.95 ) Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 04 IE TA. OYOIMPYOEY1045 PHYLOGENY POLYPODIUM AL.: ET SIGEL 2014] the P. the complexes: plesiosorum species vulgare well-supported P. two encompassing 88%,BIPP=0.95)forthemonophylyof within podium relationships evolutionary showing phylogram bootstrap likelihood (BIPP) abilities maximum con- if were (MLBS) Nodes support supported statistics). tree well for 1 sidered Table yielded (see compara- support tree and ble topology consensus identical inference of hypotheses Bayesian phylogenetic combined the the for tree and available ML are dataset best all The and 1). (Appendix generated GenBank newly in were 115 study, this in aae n osnu re r eoie nTeBS (http://treebase TreeBASE in 15263). deposited study are .org; trees consensus and dataset ihamnmmaeo 68±12mlinyasao(y;calibration (mya; ago years million 1.2 con- ± (Kvac age 26.8 Kvac Oligocene additional of E). the an age point from included minimum fossil analyses a four one Polypodiaceae with the a and of from estimate (Rothfels Two estimate straint best-age best-age 2012). the the al. of to percent et ten equal to nor- mean equal a deviation a assigned standard uncertainty with was for point distribution, compensate calibration mal To each Polypodiaceae. estimates, these the 2). surrounding for Appendix Schuettpelz (2009) from F; Pryer obtained estimates constant and A–D, best-age a points are previ- points (calibration or five calibration These constraints prior incorporated age analyses speciation derived All birth-death ously prior. a coalescence either size with population anal- run separate study species), were this single a yses intra- for of and individuals sampling species) (multiple variation across Because specific (sampling variation points. interspecific both calibration for included appropriate and is priors that model model Mkv a 2001). employ (Lewis data to variable unable ordered standard is 1.7.4 v. Beast ihetmtdprmtr n ikdte.Cddidl peetonly model (present indels clock Coded relaxed tree. linked lognormal for four a linked and The parameters 2012). a estimated al. assigned with et were Lanfear partitions 1; total) (Table data in PartitionFinder model partitions substitution by (four nucleotide determined best partition as the data Resource to separate corresponding Cluster parameters a Shared with to Duke assigned each the was for on locus data 2012) Beast Sequence al. in et (https://wiki.duke.edu/display/SCSC/DSCR). implemented (Drummond as inference 1.7.4 Bayesian v. using dataset combined the MC hoormwt endvrec ieetmtsad95% and estimates time trees divergence 32,000 intervals. (HPD) the mean density credibility posterior combine clade with highest TreeAnnotator maximum to a program chronogram used produce and The was (MCC) runs) four burn-in. 2012) for trees trees/run al. as (8,000 (20%) et discarded 2,000 (Drummond first effective were The v1.5.4 all the run 200. of when exceeded each distribution parameter sufficient of each posterior considered of the size was sample assess mixing to and used and parameters, (Rambaut and was 1.5 v. 2007) coalescence Tracer Drummond generations. size 1,000 every population sampled parameters constant (4) F. and calibra- A–D, and points coalescence A–F; calibration size calibra- population points and constant prior (3) tion F; speciation birth-death esti- A–D, (2) age points upper tion prior A–F; the speciation points birth-death in (1) calibration uncertainty were: and analyses for and four allow the 25.73 summary, and In between mya) mate. estimate (25.6 interval age fossil minimum confidence the the offset 95% approximate for closely an a settings and These in 5 mya. of resulting 44.03 mean mya, a 25.6 with distribution of exponential an assigned genus and was the sporangia, to intact belongs venation, spores visible observable with fragment frond fossilized the iegneTm Estimation— Time Divergence u hlgn Fg )poie outspot(LS= (MLBS support robust provides 2) (Fig. phylogeny Our hlgntcAnalyses— Phylogenetic oraaye eern ahepoigauiu obnto ftree of combination unique a employing each run, were analyses Four ahaayi a u ortmsfr1,0,0 eeain,with generations, 10,000,000 for times four run was analysis Each vulgare trnG-R .appalachianum P. n h soitdMB n IPspotvalues. support BIPP and MLBS associated the and eeecue rmtedvrec ieaaye because analyses time divergence the from excluded were ) ope MB 7,BP .)adthe and 1.0) = BIPP 97%, = (MLBS complex ope opie orwl-upre clades: well-supported four comprises complex ˇ k(01 rsnssrn,bticnlsv,eiec that evidence inconclusive, but strong, presents (2001) ek ru MB 0% IP=10.The 1.0). = BIPP 100%, = (MLBS group ³ ³ 0 n aeinifrnepseirprob- posterior inference Bayesian and 70% .5 nFg epeettebs ML best the present we , 2 Fig. In 0.95. ld,the clade, Results fte15DAsqecsused sequences DNA 125 the Of iegnetmswr siae from estimated were times Divergence Polypodium .glycyrrhiza P. hsclbainpoint calibration This . Polypodium ld,the clade, ˇ k2001), ek Poly- .s., s. P. .cambricum P. nlssposition analyses to oeo h hoorm nti ntne enaeestimates age mean instance, this root In designated chronogram. the the 1, of node node notably most analyses, the across than less were HPD differences individuals 95% years. 40–41), and million 0.2 36–38, time intraspecific 34, divergence 32, node uniting 30, mean 29, in 27, nodes 26, 24, 21, most (nodes million two For than time less divergence were node years. HPD mean 95% and in height) differences node 25, 39), (mean 23, 22, 35, 33, 1–20, 31, (nodes majority taxa 28, the interspecific For uniting 3). nodes Appendix the in for of found estimates be depicts (node can of analyses 2 3 the times analysis all Figure from divergence analyses. derived of four chronogram the the comparison across the tips taxa affect branch interspecific to not fos- confined differences did the primarily and Topological were of analyses E). absence the point the among (calibration to constraint or the sil coalescence) to constant or size insensitive speciation (birth-death population relatively prior model were of tree of 2012) estimations choice 1.7.4 al. v. that Beast et by suggests pre- (Drummond employed This times are divergence 3). analyses and topological four Appendix all in for sented highest (HPD) 95% similar and distribution ages and posterior identical node topologies (mean estimates nearly (MCC) time divergence produced credibility analyses clade maximum time divergence four groups three these among unsupported. relationships are The 0.95). = BIPP 70%, + Windham .aopu,P a P. amorphum, P. S. and C, G, A, of clades union among the relationships = for (MLBS support G moderate + and cladesC+S(MLBS=75%,BIPP=0.91)hintatpotential A 0.72) clades = of resolved, BIPP union poorly 60%, the 2). for are Fig. support respectively; weak clades S, but these and C, among G, A, Relationships clades as to referred nawl-upre ld MB 4,BP .) whereas 1.0), = of BIPP 94%, accessions = (MLBS all clade well-supported of a accessions Both in 1.0). = BIPP 100%, a upr MB 0% IP=10 for of 1.0) accessions maxi- two = to BIPP of provides sister 100%, S accessions = Clade (MLBS two support another. unites mal the one C to with Clade sister 1.0), unambiguously 0.58). = = BIPP BIPP 100%, 80%, cambricum = P. (MLBS support weak ifrnitdcaeof clade latter differentiated the Within 1.0). = BIPP clade, 92%, = (MLBS species four other 2), (Fig. plesiosorum decumanum 8,BP .7 o agrcaecnitn of consisting clade larger + a s. for s. 0.97) = = (MLBS BIPP support good 88%, is there phylogeny, inferred our In 1.0). Polypodium MB 7,BP .) hr ssrn upr for support strong is there 1.0), = BIPP 97%, fauriei P. = Clade Within (MLBS species. the G among differentiating for support no e oe hwdgetrvraini iegnetimes divergence in variation greater showed nodes few A Estimations— Time Divergence ld MB 0% IP=10 nldsalsmlsof samples all includes 1.0) = BIPP 100%, = (MLBS A Clade mn ebr ftemonophyletic the of members Among .colpodes P. luooipi makinoi Pleurosoriopsis .martensii P. .subpetiolatum P. ssse to sister as MB 0% IP=10,with 1.0), = BIPP 100%, = (MLBS .m + J.Sm. .s ihsrn upr MB 7,BP = BIPP 87%, = (MLBS support strong with s. s. oyoimsanctae-rosea Polypodium + and ld,adthe and clade, .rhodopleuron P. ppalachianum et ssse oawl-upre n highly and well-supported a to sister is Mett. .glycyrrhiza P. .macaronesicum P. luooipi makinoi Pleurosoriopsis loetspolypodioides Pleopeltis .californicum P. .colpodes P. .scouleri P. swl upre ssse othe to sister as supported well is , rspi contigua Prosaptia ,and MB 5,BP .) Our 1.0). = BIPP 95%, = (MLBS .scouleri P. areunitedinacladewith uz + Kunze .sibiricum P. MB 0% IP=1.0). = BIPP 100%, = (MLBS g siae rmour from estimates Age ssse pce MB = (MLBS species sister as + .californicum P. telte ihMB = MLBS with latter (the .glycyrrhiza P. L)EG nrw & Andrews E.G. (L.) .plesiosorum P. scoetsse to sister closest as .rhodopleuron P. ld (henceforth clade .plesiosorum P. .pellucidum P. ,and huhteeis there though , .cambricum P. Polypodium Phlebodium r united are MB = (MLBS group being sister + P. Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. bandb oiyn cne mgso ebru ocesue nti td.Saebr ett ahsloet ersn .4cm. 2.54 represent silhouette each to next bars Scale study. this in used vouchers herbarium of images scanned modifying by obtained the and MB)adBysa neec otro rbblte BP)aemsl ie bv oe n niae ihtikndbace seistlegend) inset (see branches thickened the with within indicated species and diploid of nodes monophyletic subclades above the major given the identify indicate mostly P S are and (BIPP) V probabilities bolded posterior The inference Bayesian and (MLBS) vulgare 06SSEAI OAY[oue39 [Volume BOTANY SYSTEMATIC 1046 with Fig. Platycerium ope,fv aablnigt the to belonging taxa five complex, .Tebs Lpyormfo h obndaayi of analysis combined the from phylogram ML best The 2. .scouleri P. h ugoptxnms itnl eae othe to related distantly most taxon outgroup the , ld,rsetvl.Tubalsloetso ebr fthe of members of silhouettes Thumbnail respectively. clade, .plesiosorum P. .vulgare P. ope,adegtotru aa(pedx1.SeTbe1frte ttsis h rei rooted is tree The statistics. tree for 1 Table See 1). (Appendix taxa outgroup eight and complex, ope n monophyletic and complex .vulgare P. .vulgare P. atpA ope:the complex: , rbcL ope codn oShedre l 20) Lbosrpspotvalues support bootstrap ML (2004). al. et Schneider to according complex , matK, .appalachianum P. .vulgare P. and .plesiosorum P. trnG-trnR complex, ld,the clade, eunedt o e ili aao the of taxa diploid ten for data sequence ope,rsetvl.Ble etr ,G ,and C, G, A, letters Bolded respectively. complex, .plesiosorum P. .glycyrrhiza P. ,and ld,the clade, luooipi makinoi Pleurosoriopsis .cambricum P. Polypodium clade, were . Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 04 IE TA. OYOIMPYOEY1047 PHYLOGENY POLYPODIUM AL.: ET SIGEL 2014] niae nFg .Bakcrlswt etr niaetesxclbainpit sn nti td:pit – n eeotie rmSchuettpel from obtained were F and Kvac A–D points by study: reported this fossil in a using to points calibration refers six E the point indicate and letters (2009) with circles Pryer Black 2. Fig. in indicated oeb h rybr.Telwrlmto h 5 P o oe1i rnae.Dtsfrgooi eid n pcsa ersne ntebto bar bottom the in represented as epochs and periods each geologic for for indicated Dates are truncated. HPD is 95% and 1 age node Mean for 3. HPD Appendix 95% in the (2009). reported of Geissman are and limit node Walker lower each to The to correspond bars. corresponding grey Statistics the 41). by (node node youngest to 1) (node oldest Fig. .Mxmmcaeceiiiy(C)crnga ihtedvrec ieetmtsfo nlss2 ldsP ,A ,C n r as are S and C, G, A, V, P, Clades 2. analysis from estimates time divergence the with chronogram (MCC) credibility clade Maximum 3. ˇ k(01 pedx2.Salnmesaoendsidct ennd g siae from estimates age node mean indicate nodes above numbers Small 2). Appendix (2001; ek zand Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 08SSEAI OAY[oue39 [Volume BOTANY SYSTEMATIC 1048 oeb h lgcn osldtda 68 . y (Kvac mya 1.2 that ± 26.80 on at imposed dated constraint it fossil Oligocene age Thus, the minimum by the 4). node reflect and ranged 3 and 2 analyses constraint for 1 (analyses times divergence estimated age mya older the that 24.77 fossil appears whereas and the respectively), 24.65 from 3, incorporating between ranged and node not 1 that those (analyses for mya constraint age 27.64–27.80 fossil a porating makinoi. Pleurosoriopsis ta.20;Ot ta.20) h ngai sa taxon Asian enigmatic the 2009), al. of groups et (Schneider Otto makinoi studies Haufler Pleurosoriopsis 2004; molecular previous 1982; al. in As Tryon et b). and a, 1995 Neotropical Tryon al. et 1928; the (Christensen with group relationship sister its 2.46– HPD: (95% HPD: mya (95% 5.50 respectively. mya and 16), 3.72 node 17) mya; 14), 8.93 node at mya node mya; 8.81 estimated mya; 20), is 1.53–6.24 node 5.06–13.08 mya; S HPD: 0.86–3.85 and HPD: (95% C, (95% 12). mya G, node 2.18 mya; A, about 7.45–16.76 clades HPD: within (95% mya Diversification 11.91 and 10) node 13.60 approximately majo between the the into Within divergence 13). plex, mya node 11.38 started 6.90–16.57; HPD: have (95% to estimated is diversification monophyletic group, to reciprocally leading the and 8), complex node of mya; formation Clado- 14.01–27.06 7). the HPD: (95% node mya mya; within 20.59 16.72–32.74 events HPD: genetic (95% mya 24.77 and s. s. h otrcn omnacso of ancestor common recent most within the dates divergence conclu- age our and an affect Polypodium not relationships placing does regarding it not but sions from node, 3). root results (analysis the on mya variation constraint 99.33 this to 2) suspect (analysis We mya 78.57 from range lsi oi(i.2 togyspottemnpyyo the of monophyly the support north-temperate strongly primarily 2) (Fig. loci plastid 2006). al. et Smith 2006; al. et and such genera Sm., segregate of series a as the of recognition for the of evidence supporting circumscription molecular narrowed this compelling even of provided polyphyly Otto (2009) and 2006) al. (2004, al. et 1995b; et al. Schneider et 2004). Smith Haufler and 1995; Mickel Moran spe- 1982; World century Tryon New and 20th (Tryon primarily cies late 100–150 approximately many name of 1973 generic and group the Sota of 1990), la application limit al. De authors et (see has years Hennipman circumscription the and This over the sori. significantly dis- (1753), narrowed with round ferns Linnaeus non-marginal, leptosporangiate by crete, most described encompassed originally genus As history. analyses. four all the among in estimates overlap substantial HPD with 95% Oligocene, divergence late the mean in 7 a node indicate for analyses all Nevertheless, 2001). h iiuie ihydvddlae Fg ) prni fol- sporangia 2), (Fig. leaves divided highly diminutive, the u hlgntcaaye fsqec aafo four from data sequence of analyses phylogenetic Our of taxonomy The that infer we 3), (Fig. 2 analysis on Based Pecluma Polypodium. Synammia luooipi makinoi Pleurosoriopsis Price, .s rae aito a lose o oe7, node for seen also was variation Greater s. s. .plesiosorum P. Phlebodium Polypodium .Pel(afe ta.19;Schneider 1993; al. et (Haufler Presl C. hsrmisasrrsn eutbecause result surprising a remains This endvrec ieetmtsincor- estimates time divergence Mean sifre ob itrt hs two these to sister be to inferred is Discussion Polypodium oyoimvulgare Polypodium ru.Wti the Within group. (R.Br.)J.Sm., y 9%HD .81.7mya; 8.98–18.67 HPD: (95% mya rcladesA,G,C,andSoccurred a ogadcomplicated and long a has iegdapproximately diverged ea approximately began Polypodium Serpocaulon .vulgare P. Polypodium .plesiosorum P. .plesiosorum P. ope and complex Polypodium Polypodium .vulgare P. .s and s. s. .R. A. com- oa to ˇ ek , ula eunedt a eov hs pce relationships species these resolve incorporating may data studies arctic/ sequence nuclear an molecular and Further sporangiasters distribution. eglandular its boreal by these of both otenCsaeMutiso rgn ahntn and Washington, glandular) Oregon, Columbia. of still British Mountains (but Cascade few northern >52 relatively spores and pinnae, sporangiasters, obtuse short, Canada. eastern to and Appalachians sporangiasters, glandular abundant spores b). apices, a, 1995 acute al. with et Haufler 1993; appalachianum al. Polypodium and et (Haufler Haufler morphological distribution 1991; cryptic, geographic Windham distinct somewhat a if and stable, features a has of taxon suite Each species. unique circumscribed broadly single a into eainhp mn aai ld ,w ontadvocate not do we A, clade in collapsing genomes taxa among plastid relationships and the loci) between be isozyme evolution al.1987). might et the of (Wolfe discrepancy (coding rates This differential nuclear relatives. dis- genetically by close is each explained its that from suggesting within (aver- 0.46), taxa tinct vs. values between 0.96 those identity of than genetic higher age much that were taxa reported these of (1995b) Haufler light al. group. in the et of surprising studies is electrophoresis resolution isozyme earlier interspecific of lack This speci- included All of 2). mens (Fig. species individual of monophyly n niiulof individual one assalsmlsasge otrepiaiyNrhAmericantaxa: North primarily three to assigned samples all passes MB 7)adpoie ekspot(LS=6% for 62%) = of (MLBS clade support of combined weak a provides individuals of and 87%) sampled exception = (MLBS three the the with unites indi- datasets, from locus of derived topologies lack vidual across This consistent differentiation. is sequence resolution little with polytomy a ( dataset support combined no the recovers for topology likelihood morphological maximum unambiguous best an provides ii) group. sporangiasters and the of for synapomorphy presence i the previous al. and 1 2009), et all al. (Fig. Haufler et in Otto 1995; recovered b; Ranker a, been 1995 and (Haufler has studies Strong clade molecular 1991). this Windham 1969, for and (Lang support Haufler Lloyd revisions 1974; Siplivinski taxonomic of 1971; subsequent members “ by diploid expanded (1964) the Lang’s to and correspond taxa These isi ih fhptee rpsdb rvosresearchers. previous by proposed spe- hypotheses these of among light relationships in cies inferred the discuss we Here oee,i ospoiefracnein n unequivocal and of to sister convenient clade 2013) monophyletic a Haufler for of provide demarcation and does Shugang it 1995; However, al. Ikebe et Pleurosoriopsis and (Kurita Iwatsuki spores chlorophyllous 1977; and veins, the lowing togspotfrfu ldso ili pce belonging species diploid of the clades to four for Complex— support strong vulgare Polypodium the ept h nblt fpatdsqec aat resolve to data sequence plastid of inability the Despite ept togspotfrtemnpyyo ld ,our A, clade of monophyly the for support strong Despite P hlgntcRltosisAogDpodMmesof Members Diploid Among Relationships Phylogenetic LPDU APPALACHIANUM OLYPODIUM .aplcinm .amorphum, P. appalachianum, P. .vulgare P. < .appalachianum P. 52 .appalachianum P. m og t ag xed rmtesouthern the from extends range its long; m r odfeetfo hs ftypical of those from different so are Polypodium ope Fg.2 ;cae ,G ,adS). and C, G, A, clades 3; 2, (Figs. complex oyoimsibiricum Polypodium .sibiricum P. .appalachianum P. oyoimvirginianum Polypodium < 0 LSand MLBS 50% , .amorphum P. a eaieyln,nro pinnae narrow long, relatively has .s,wihcnb eie sthe as defined be can which s., s. luooipi makinoi Pleurosoriopsis , CLADE .amorphum, P. m ( matK og ti ofndt the to confined is it long; m oyoimamorphum Polypodium A—hscaeencom- clade (A)—This and + u nlssprovide analyses Our oooyntshown). not topology and , sdsigihdfrom distinguished is .amorphum P. < .sibiricum P. .0BP)frthe for BIPP) 0.70 .appalachianum P. .sibiricum P. and .gop,as group”, L. matK .sibiricum P. . Polypodium itrto sister Fg 2). (Fig. which form has . Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. h apigfrti td) oprdto compared study), in this variation for molecular sampling greater the to due = BIPP glycyrrhiza be P. and could 80% = This MLBS 0.58). 2, of (Fig. monophyletic rachises as supported the whereas contrast, clade, of individuals well-supported sampled In two The 1995). 1993). al. glycyrrhiza (Christensen et P. hairs rachis Iwatsuki adaxial gray 1928; long, the and dried of when members diploid vulgare other P. Unlike morphology. other unique in found glycyrrhiza not P. trait that a 1995), indicate the al. of et members Iwatsuki diploid 1993; Haufler al. 1928; (Christensen leaves et the of surface hairs adaxial multicellular the of on presence the by united morphologically of and relationship sister the for support “ unequivocal the of members diploid the sister species infer 1995a; al. diploid to 2009). et al. ability et three Haufler Otto 1995; the the Ranker limiting and of (Haufler group, relationships two this than to belonging more the no within included relationships addressing including and clade a to sister being subsequent ( support A weak 1995b). utilizing al. et study Haufler Peninsula; Kamchatka fauriei supported tributions 04 IE TA. OYOIMPYOEY1049 PHYLOGENY POLYPODIUM AL.: ET SIGEL 2014] eandursle Fg ,b.Smlrte nisozyme that in to suggested sister Similarities be species b). might profiles these a, banding 1 among electrophoresis (Fig. relationships unresolved However, remained group. with species this Asian identity the that genetic sized of levels glycyrrhiza high isozyme indicating on Based studies America. North to western confined in largely habitats are coastal and sporangiasters, lack surfaces, leaf yohszdacoerltosi between relationship and close a hypothesized Asia: eastern californicum, Rim and Pacific America the North occupying western species of three of samples all passes sad smaie nRme ta.21) Alternatively, 2014). al. et Rumsey in with been (summarized allied, Islands long closely has are cambricum taxa It treating two authors sporangia. some these the that among Wagner hypothesized 1950; Martens scattered iv; and (distinctive iii 1964) 1 paraphyses Fig. see branched hairs; receptacular having by characterized macaronesicum of accessions two otenClfri Hulre l 93 adi ta.2012; and al. central et Baldwin 2014). to al. 1993; et limited al. Sigel et largely (Haufler is California southern range geographic whose within evolution morphological of clade. inferences this for allow and u td rvdssrn upr o h oohl of monophyly the for support strong provides study Our P P LPDU CAMBRICUM OLYPODIUM GLYCYRRHIZA OLYPODIUM .scouleri P. .glycyrrhiza P. .glycyrrhiza P. secuieyAinand Asian exclusively is ra useisrsrce oteMacaronesian the to restricted subspecies a as or , complex, and and + and cosalre egahcrne(srfetdin reflected (as range geographic larger a across Fg ;Hulre l 00.Alohrstudies other All 2000). al. et Haufler e; 1 (Fig. Fg ld )it elspotdclade well-supported a into C) clade 2 (Fig. .californicum P. < .californicum P. .fauriei P. trnL-trnF 0 aiu asmn S for BS) parsimony maximum 50% .californicum P. .californicum, P. .fauriei P. eas hyhv ar nteradaxial their on hairs have they because .cambricum P. Fg ld ) hs he pce are species three These G). clade 2 (Fig. .fauriei P. .fauriei P. .macaronesicum P. CLADE .vulgare P. lsi eunedt provided data sequence plastid ssrnl ifrnitdfrom differentiated strongly is Fg ) ly n ag(1964) Lang and Lloyd 2). (Fig. CLADE .glycyrrhiza P. .fauriei P. afe ta.(95)hypothe- (1995a) al. et Haufler , Fg ) atr elce nits in reflected pattern a 2), (Fig. a itntv uvdfrond curved distinctive a has and .glycyrrhiza P. C—u nlssuie the unites analysis (C)—Our .glycyrrhiza P. r ueuet(afe tal. et (Haufler puberulent are .gyyria .californicum P. glycyrrhiza, P. n u nysml of sample only our and hra igorpi dis- biogeographic whereas G—hscaeencom- clade (G)—This .glycyrrhiza P. ope.Bac lengths Branch complex. a loamme of member a also was .californicum P. .vulgare P. sasnnmof synonym a as ru”adnovel, and group” .gyyria P. glycyrrhiza, P. xed nothe into extends sol weakly only is .californicum P. .californicum P. .californicum P. ssse ( sister as .fauriei P. .fauriei P. complex oma form P. P. P. P. , .pellucidum P. recovered Fg ) ae td using study later A d). 1 (Fig. resolve to surprised were n ag16;Hulre l19 ,b). a, (Lloyd al.1995 speciation et reticulate Haufler in 1964; part Lang no and or little played have to widespread the of to studies netic pointed who across Nevertheless, (1981), venation anastomosing Lellinger of homoplasy by part, venation, in of anastomosing species into goniophlebioid strongly Neotropical its Polypodium the that with with suggested belonged species, (1928) Christensen 1999; this debate. Haufler historical to and much endemic Li is 1997; and Li 2003). veins Palmer 1888; free (Hillebrand (pellucid) Hawaii Mickel 2012). transparent 1993; al. al. et has Baldwin et 2004; (Haufler Smith America and hab- North coastal western to in restricted f). itats is by 1 and recovered veins Fig. anastomosing was (2009; strongly relationship al. of sister et texture their Otto leathery with thick, and the associated leaves in been their similar rarely are having they another, Despite one S). clade 2 clade, (Fig. well-supported this in o upr o,rltosisaogtefu ili clades. diploid four the among relationships for, support low the of phylogeny cnrowudepanwypeiu tde resolved studies a Such previous 2000). why Raquin and scouleri explain P. Guillon either would 1998; al. from scenario et genomes Vogel 1992; plastid californicum inherited P. Marin maternally they, that in possible with collected is it were and resembling specifically, despite studies Reyes Pt. molecular and County earlier of the individuals The in 1951; 2002). (Manton California al. Hill, et Tank Hildebrand and Reyes Pt. from 1991) from hybrid triploid a be of may between recognition studies subsequent earlier the of by those explained and results our between ancy with ..Rme,Crn ob ra itntspecies, distinct a as or Robba & azoricum Carine Rumsey, F.J. as Islands of Azore the populations recognize authors some hc ute uprstesgeainof segregation from respectively), of the England, supports monophyly and further Spain the which 2; al. Fig. confirm 1; et (Appendix to Rumsey range taxa able 2009; are two al. cambricum et also these Otto We 1995; 2014). between Ranker and divergence (Haufler genetic showed that significant analyses phylogenetic previous reinforces study recognition abun- the support and that of shape structure scale paraphysis rhizome and serration, dance, differ- lamina documented (1980) in Roberts ences 2014). al. et Rumsey 2005; afe n akr(95 nlddboth included (1995) Ranker and Haufler of placement taxonomic The R P .cambricum P. LPDU SCOULERI OLYPODIUM LTOSISAOGCLADES AMONG ELATIONSHIPS Serpocaulon .californicum P. .cambricum P. .californicum P. .scouleri P. Vs. o.Frads(afe ta.20;Scha 2000; al. et (Haufler Fernandes Ros. (Vasc.) ssse to sister as .scouleri P. ayo hc aebe eetysegregated recently been have which of many , rmtesuhr n otenprin fits of portions northern and southern the from ntefirst the in .scouleri P. or and Sihe l 06.Ti dawsdispelled, was idea This 2006). al. et (Smith . .vulgare P. .glycyrrhiza P. .vulgare P. n h allotetraploid the and Fg ;Hulre l 00.Tediscrep- The 2000). al. et Haufler e; 1 (Fig. .scouleri P. .macaronesicum P. and .macaronesicum P. and .glycyrrhiza P. CLADE a xlddfo eea phyloge- several from excluded was .glycyrrhiza P. rbcL .glycyrrhiza P. .scouleri P. oyoimscouleri Polypodium ope eas twsthought was it because complex ope it t u provides but at, hints complex eeatal rpodhybrids triploid actually were , S—w pce r included are species (S)—Two trnL-trnF hlgn of phylogeny ,G C, G, A, .scouleri P. .scouleri P. GsoyadYatskievych and (Gastony ,ratherthan ssse to sister as sdsic pce.Our species. distinct as subsp. ;WhitmoreandSmith oyoimpellucidum Polypodium lsi eunedata sequence plastid ntdi polytomy a in united AND .macaronesicum P. a enamte of matter a been has .calirhiza P. .scouleri P. and .macaronesicum P. —u inferred S—Our Polypodium azoricum Polypodium .scouleri P. .pellucidum P. .pellucidum P. .glycyrrhiza P. a obscure, has included (derived (Vasc.) and , and ¨ .l. s. fer on P. P. . Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 00SSEAI OAY[oue39 [Volume BOTANY SYSTEMATIC 1050 vulgare P uigteeryMoee(i.3nd ;2. y,95% mya, restrial 20.6 of epiphytic 8; divergence mostly the node for the estimates 3 These (Fig. mya). Miocene 14.00–27.06 HPD: early the during (P) diver- subsequent the the of and gence mya), 16.72–33.52 Oligocene/ HPD: late 95% mya, a diver- of for origin Our support Miocene 2013). early provides Haufler analysis and time Shugang gence 1993; 1964; al. (Valentine America et North Haufler and Asia, Europe, of temperate northern regions the in exceptions, the few of with members occur, complex contrast, In 2012). Central al. Hispan through et and (Luna-Vega America, Mexico South from central Cancer, and northern of America, Tropic the of south subpetiolatum P. the to assigned formerly species (including group the of Members in non-overlapping. largely clades are monophyletic s. s. large two the of tribution w pce and species of those two to similar most were describ- scales In clades. two these to ing affini- belonging taxa suggested the have among ties authors in previous Basin some Pacific and is distribution), C is S C clade clade (clade whereas between separation European/Macaronesian, geographic relationship broad and sister their Despite 75% the S. = = clade is BIPP MLBS 0.91) and 2; = 60% (Fig. = BIPP supported MLBS are better 2; Somewhat American, (Fig. 0.72). North support predominantly weak both with united G, and A Clades oentoia anfrss(cutpl n re 2009). Pryer and of (Schuettpelz rise forests rain the tropical followed modern that ferns epiphytic of radiation Cenozoic pcieo h rgno h diploids. per- the new our of origin provide on al. the to on spective able estimates et are (Haufler date we taxa phylogeny, divergence “diploids-only” diploid calculating the By of 2000). speciation and clad- precipitated ogenesis have may that events historical the been accorded has climatic attention Less changes. and distributional these invoked explain geographic been to have glaciation, Major Pleistocene the past. as such geo- the changes, had in present, that at contact indicate allopatric graphic be species may allopolyploid which species, existence, 2005). diploid Yatskievych very and their Windham Zhongren 1995b; By and al. (Haufler et taxa Haufler allotetraploid 1991; the of origins the diploid the the regarding hypotheses of the evolutionary address- distributions of the members for geographic shaped have and framework that relationships events new historical a the ing provides phylogeny the Our of Members Diploid .cambricum P. of developingsori the the noted in paraphyses (1997) branched deciduous Li of addition, presence In between (0–0.79%). crosses congeners diploid to relative and (5.95%) between C crosses successful clade of percentage of higher a demonstrated members He observations. phological the pellucidum between P. affinity sug- also an (1997) Li gested climates. oceanic with regions unglaciated P P Among Phylogeography and Estimates Time Divergence LPDU VULGARE OLYPODIUM OLYPODIUM .macaronesicum P. ope V,a nta iesfcto curdduring occurred diversification initial an (V), complex .vulgare P. .pellucidum P. .vulgare P. and ae nhbiiaineprmnsadmor- and experiments hybridization on based ES STRICTO SENSU rus cu oeyi h etois mainly Neotropics, the in solely occur groups) .vulgare P. .macaronesicum. P. .cambricum P. ope ogl orsodt h mid- the to correspond roughly complex hc r iia otoefudin found those to similar are which , .plesiosorum P. orv(94 oe htisrhizome its that noted (1964) Bobrov , ope V and (V) complex Polypodium COMPLEX .vulgare P. Tepeetdygorpi dis- geographic day present —The oyoimvulgare Polypodium ru.T ae phylogeographic date, To group. .pellucidum P. r l on npersistently in found all are Wti the —Within ru n h agl ter- largely the and group ope aefcsdon focused have complex .s Fg oe7 24.77 7; node 3 (Fig. s. s. .scouleri P. .pellucidum P. .plesiosorum P. and .macaronesicum P. n htthese that and , .plesiosorum P. .dulce P. Polypodium Polypodium Complex— n other and .vulgare P. group ˜ and ola ot mrc saoe40 above is America North AieadWle 98 oe n aeet19;Willis 1998; lineages Kadereit animal and of Comes and 1998; mechanism plant Walker a other and as (Avise numerous invoked in been speciation has refugia likely Pleistocene are and 2007; amorphum 2009) Paris of P. al. speciation and allopatric et the and Barrington Walker for Steele locations 2006; 2008; 1997; Beebee al. and al. et Zeisset et Soltis Soltis 2006; and the 1958; Storfer plant in (Detling many for species Valley refugia animal Pleistocene River known Columbia well are and west Peninsula and Olympic east during the refugia the in Mountains south coastal Appalachian or shifted The advances. montane range glacial to its restricted but became Pleistocene, and the to prior Canada and r,tebgnigo eetdcce fgaito (Walker glaciation of cycles repeated 2009). bound- mya). Geissman of and Pliocene-Pleistocene 0.86–3.85 beginning HPD: the of the 95% to mya, ary, group closely 2.18 crown 20; corresponds node the This 3 for (Fig. relatively A date diver- a clade providing divergence little and estimated showing analyzed loci recent by plastid hypothesis the study at this Our gence America. pre- to North speciation credence in allopatric lends events that glaciation from hypothesize by resulted to cipitated taxa (2000) diploid al. distri- these et geographic Haufler Their prompted 1993). al. butions and et (Haufler Haufler Alabama 1991; to Windham south Mountains Appalachian along 1969; Newfoundland the from (Lang extending America Valley 1993). North River eastern al. Columbia et the Haufler along north- and Oregon Washington, ern in Mountains Olympic and distributions. Cascade southern and amorphum limited and more Shugang 1993; have al. et 2013). (Haufler Haufler northern Siberia Canada, and western China, and Japan, central in distribution boreal n rs 97 ilu19;Hulre l 00.W hypothe- We of 2000). ancestor common (Dyke last al. the et that ago the Haufler size 1991; years during Pielou thousand 1987; Sheet Prest 21–18 and Ice (LGM), Laurentide maximum the glacial of edge southern eierna anaddsrbto,adoeedmcto or maritime endemic primarily one a islands. and volcanic with distribution, one species, mainland two Mediterranean and in only C occurring clades have taxon both a 2) S taxa; of American composed North are with together G Asia and A clades: clades boundary. resulting both the Pliocene-Pleistocene 1) among emerge the patterns geographic American Two near North diversifying the of taxa majority late the the until begin with not did Miocene-Pliocene, clades species major extant four the of of each diversification within Subsequent 2009). Price (Rundell and opportunities con- ecological environmental novel 2008). changing diversifica- provided by that of ditions Kjer driven burst was this and likely radiations most unclear, tion Whitfield are rapid specifics 2007; the ancient Although Lockhart topo- of and approxi- a of characteristic show (Whitfield span divergences pattern a These 3 within years. logical (Fig. diverging million S) clades two and extant mately C, major G, four A, all clades with mid-Miocene, the oet fNrhAeiaadesenAi,ec ihisown temperate its range. with the each geographic in Asia, eastern distinct outcrops and rocky America North inhabit of forests A clade in species oto h rsn-a ag of range present-day the of Most P LPDU APPALACHIANUM OLYPODIUM .amorphum P. scnie osuhr rts ouba the Columbia, British southern to confined is epciey egahcioaini multiple in isolation Geographic respectively. , oyoimappalachianum Polypodium a ral itiue cosboreal across distributed broadly was oyoimappalachianum Polypodium oyoimsibiricum Polypodium  CLADE aiue hc ak the marks which latitude, N .sibiricum P. A—h he diploid three (A)—The oyoimsibiricum Polypodium .appalachianum P. and , .appalachianum, P. .amorphum P. sfudin found is a wide a has Polypodium and in Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. 04 IE TA. OYOIMPYOEY1051 PHYLOGENY POLYPODIUM AL.: ET SIGEL 2014] eodr otc nteSnFacsocatlbsn(Baldwin these 2012). basin al. coastal et Francisco Maximum, San the Glacial into in coming contact ranges, secondary Last present their into the expanded likely species Following 2004). 2001, allopatric the of reinforced speciation and triggered have may glaciation of cycles repeated during refugia in southern Isolation and northern Northwest. separate Pacific common the through the Alaska Pleistocene, from range the 1981; to of (Lindsay prior ancestor 2005). Oberbauer that 2001; advances hypothesize al. et We Maldonado glacial 2001; al. Pleistocene et Arbogast 2004; species the northern harbor Smith to of thought during and also areas range (Mickel 2014), al. present et Island Sigel Guadalupe The and 1997). Valley peninsula al. River et Columbia californicum Soltis the 1991; and (Pielou Islands, the Island, Charlotte Vancouver Queen including refugia, coastal Pleistocene of range current The Pleistocene. geographic the of cycles climatic and the precipitated by was reinforced that sug- speciation allopatric distributions, of geographic history a their gest with together mya), 4.12 californicum P. P. between divergence Pliocene-Pleistocene tdjs otes fteKrlIlns ak h western the marks Islands, Kuril the of of 1995). of extent Islands situ- northeast al. Peninsula, Kuril just Kamchatka the The et ated 1995). and al. Iwatsuki et Okinawa, (Iwatsuki except Russia 1993; Japan of al. and islands (Whitmore et California fauriei Haufler Polypodium Baja the to 1991; along Japan arc Smith continuous from nearly Rim a in Pacific distributed are G clade nnrhatr saadnrhetr ot America North northwestern and forming Asia (Hulte were northeastern compositions boreal species and in submerged dissimilar was with Bridge Land forests Bering the when corresponds and to of lineages, American establishment North the and marks Asian 5.06– distinct time HPD: divergence 95% mya, This 8.81 mya). 14; node 13.08 3 (Fig. Miocene Late the in 1993; 2014). al. al. et et Sigel (Haufler 2012; peninsula al. et California Baldwin Baja the Bay into coast Francisco of San limit The of ern limit 1993). southern the al. the of marks et area mainland Northwest (Haufler Pacific of States the coast United and the Columbia, along British south Alaska, continues then Islands, and u ok erteMdtraen u trne sfar as ranges it but Mediterranean, the near rocks ous bution. vulgare P. ocdgntcdfeetainaogteetretx (Hewitt taxa 2001). three 1996, rein- shaped these likely, glaciation among and, differentiation of America genetic cycles forced North that in evident mya; distribution is 3.8 current it as their 20), early node (as 3 of glaciation Fig. speciation Pleistocene the the of out southern beginning rule and/or to impossible appalachianum Beringia, P. is Asia, it from While refugia. dispersal by gla- range the As 2004). retreated, Hewitt ciers 2001; Knowles 2000; Whittaker and P oyoimfauriei Polypodium P LPDU CAMBRICUM OLYPODIUM GLYCYRRHIZA OLYPODIUM .macaronesicum P. ´ 98.Smlrt h pce ncaeA h estimated the A, clade in species the to Similar 1968). n oyoimcambricum Polypodium .glycyrrhiza P. ope ohv uoenMcrnsa distri- European-Macaronesian a have to complex .glycyrrhiza P. nldstePcfccato h aaCalifornia Baja the of coast Pacific the includes .californicum, P. .glycyrrhiza P. Fg oe1;25 y,9%HD 1.14– HPD: 95% mya, 2.52 18; node 3 (Fig. .sibiricum P. and cuso hj sado oe,almajor all Korea, of Island Cheju on occurs iegdfrom diverged .amorphum P. .glycyrrhiza P. hsseisac costeAleutian the across arcs species This . r h nydpodmmeso the of members diploid only the are and CLADE CLADE and ieyepne noispresent its into expanded likely hc xed ot ln the along south extends which .californicum P. sms bnato calcare- on abundant most is .californicum P. G—h he ebr of members three (G)—The .glycyrrhiza P. (C)— .glycyrrhiza P. curn utpirt the to prior just occurring nopse numerous encompasses oyoimcambricum Polypodium pne coastal a spanned glycyrrhiza + n h north- the and Hwt 1996, (Hewitt .californicum P. and P. uto fa ppyi netrt aai olwdby followed Hawaii, to ancestor epiphytic an of intro- duction single from a 1999; hypothesized derived (1997) Haufler estimates Li electrophoresis, and distance isozyme genetic (Li Using cinder 2003). and Palmer flows lava barren inhabits rainforests. in contrast, trunks tree In on living epiphyte an as exclusively n itnthbtt L 97 iadHulr19;Palmer 1999; Haufler and Li 1997; (Li 2003). habitats distinct occupy- ing varieties morphological described four with endemic 02.Teohrseiso ld is S clade al. of Baja et species Baldwin other of 1993; The west 2012). al. et Island Haufler Guadalupe 2005; (Oberbauer reported on California population sites disjunct elevation a high with from area, the to Bay high Columbia Francisco and British San zones southern coastal from salt-spray ranges the coastal rainfall to on Bailey restricted epiphyte and is (Sillett an It trees as 2003). coniferous of or trunks rocks and branches on the growing America, North or ern oceanic temperate habitats. in forest found foggy primarily taxa of composed is arslahbttmyhv otiue oteeouinof evolution the to contributed macaronesicum P. have may Emerson habitat ancestral the 1987; laurisilva of fragmentation Mai and divergence 1979; (insular) patric (Sunding Mediterranean Basins Rodrı the 2002; Sea only Black in persisting and patches Europe, isolated continental 3 to on with corresponds (Fig. laurisilva mya) diminished Pliocene greatly 1.53–6.24 of the time HPD: a in 95% mya, species 3.72 laurisilva. two 17; Tertiary node these the of within divergence distributed The broadly was and north- and Rodrı Walker and 2009; mya; Europe Geissman (2.6–65.5 were Tertiary southern the that during in Africa forests diverse western evergreen and subtropical extensive humid, most a Mediterranean is the (laurisilva), of forest the laurel relic the in islands, Macaronesian found the type vegetation dominant of group The 2006). Carine 2004; sister al. et (Carine a plant have Macaronesian of species 75% approximately whereby pattern, distribution floristic Mediterranean-Atlantic well-established Canary Scha 2007). the 1994; al. and et Short Azores, and the (Press Madeira, Islands in 1997). epiphyte Page 1964; common (Valentine Isles macaronesicum British podium the as north aie nVnepotne l 2007). al. (sum- et Vanderpoorten lineages American in within bryophytes, marized North nested temperate Macaronesian phylogenetically and and are Numerous Neotropical C ferns explain 3). clades and help and lycophytes, between 2 would relationship (Figs. and taxa S sister 2005), American apparent of al. spread the et the archipelagos (Sim-Sim for stepping-stone Macaronesian eastward a previously the expanded as a that acted with then consistent hypothesis and is proposed scenario Mediterranean a the Such northward. in evolved Europe true, to If likely event dispersal Africa. dispersal subsequent North with distance or America, long North a from following Macaronesia on icn Fg oe1;1.2ma 5 P:7.45– HPD: 95% mya, 11.92 ancestor late common 12; recent to of most middle the node that the suggesting mya), 3 during 16.76 diverged (Fig. have to Miocene estimated is C oyoimcambricum Polypodium P that possible is it Alternatively, .cambricum P. LPDU SCOULERI OLYPODIUM oyoimpellucidum Polypodium ´ guez-Sa .pellucidum P. and and ´ ce n roo21) hs ohallo- both Thus, 2011). Arroyo and nchez ´ .macaronesicum P. .cambricum P. guez-Sa oyoimscouleri Polypodium sedmct aaoei hr ti a is it where Macaronesia to endemic is CLADE and oyoimcambricum Polypodium var. ´ S—uhlk ld ,caeS clade C, clade like (S)—Much ce n roo21) Clade 2011). Arroyo and nchez .macaronesicum P. var. vulcanicum . .macaronesicum P. ¨ pellucidum .pellucidum P. e 05 Vanderpoorten 2005; fer a aeeovdin evolved have may sedmct west- to endemic is stretiland terrestrial is cusalmost occurs ofr oa to conform Hawaiian a , ol have would evolved Poly- Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. SihadTejero-Dı and (Smith bination rnfre otegenus the to pre- transferred hypotheses biogeographic other required the and herein. be by sented this will unaffected address data relatively genetic to was population Northwest, Additional distribution Pacific LGM. its the of that that areas and (Oberbauer possible coastal unglaciated equally advances in is sisted glacial it Pleistocene However, during 2005). of displacement southward flora Island the Guadalupe during the refugium of a highlands as northern rela-served the foggy that Baja cool, possible the is tively and It California 1965). of (Raven west peninsula California islands coastal the in distri- disjunct butions with species plant northwestern of assemblage of distribution refugia. geographic Pleistocene involving day present the scouleri P. G, and of A distribution clades origin, or the explain morphology, to unique hypothesis alternative an evoke readily glycyrrhiza P. the west that the by to 2011). aided Clark Pacific and likely the (Gillespie America in America, North of latitudes coast North tropical of from jetstream to coast Hawaii from Pacific from dispersed the have dispersal would scenario, ancestor following this common Under the perhaps Macaronesia. or origin, Europe Mediterranean Hawaiian of 2007; is pos- of al. ancestor the common exclude the et Vernon cannot that 2010; sibility we Geiger Alternatively, Wagner 2013). and hypothesis 2003; Ranker Baldwin and this al. 2007; al. et to et Nagalingum Ranker support 1993; lends (Barrington angiosperms Hawaiian endemic and several of ferns of origins American habit North the epiphytic for eight shared the of The scouleri all Islands. to Hawaiian events Miocene, late main dispersal the subsequent during by America North followed of Coast Pacific with the ancestor We common a 2002). of dispersal Clague distance current and the (Price of that Islands hypothesize oldest corre- Hawaiian the closely Kauai, main mya) of eight emergence 2.46–8.93 the HPD: to 95% sponds mya, 5.50 16; node ogdsac ipra i ido ae.Teestimated The water. or between time wind by divergence immigrated via have dispersal larger must a long-distance to biota connected non-endemic been all never have of landmass, and 2007) 1994; ancestor al. Dalrymple et and the Geiger Clague mya; for 80–0.5 origin (approximately gin American North pellucidum a of bility o hspoet h ..Ntoa oetSrieRgo idygranted specimens kindly collected 6 Region who Service Vernon Forest collections, L. National live A. U.S. their and The project. to Huiet, this L. access for Many Ebihara, allowing extraction. A. for DNA to RBGE collections for and and their material UCBG lending leaf of in to removal generosity thanks the their allowing for for UC and and E, DUKE, ALA, 02SSEAI OAY[oue39 [Volume BOTANY SYSTEMATIC between ancestor this established. of not provenance was Haufler geographic and the (Li However, habitats 1999). volcanic to adaptations subsequent 1052 oeaddi proof: in added Note hypothesis previous a contradict study this of results The u hlgn hw elspotdsse relationship sister well-supported a shows phylogeny Our Acknowledgments. .scouleri P. and loei sanctae-rosae Pleopelis .scouleri P. eas h aainIlnsaeo ocncori- volcanic of are Islands Hawaiian the Because . nNrhAeiacudvr elrfetahistory a reflect well very could America North in .pellucidum P. vle rmamrtm-dpe ouainof population maritime-adapted a from evolved HulradRne 95,btte onot do they but 1995), Ranker and (Haufler .pellucidum P. ´ and z2014). ez etaktehraimcrtr n tf at staff and curators herbarium the thank We Pleopeltis .pellucidum P. var. .pellucidum P. oyoimsanctae-rosae Polypodium oyoimscouleri Polypodium pellucidum rgntdfo ige long single, a from originated Mxn .R m Tejero & Sm. R. A. (Maxon) n sindtenwcom- new the assigned and .pellucidum P. ugsigtepossi- the suggesting , and swl sevidence as well as , .scouleri P. .scouleri P. .scouleri P. .scouleri P. and spr fan of part is .Aswith a been has .scouleri P. Fg 3 (Fig. from per- P. P. ek .B,M .Wnhm .Ytkeyh n .M re.21.A 2010. Pryer. M. K. and Yatskievych, G. Windham, D. M. B., and Rosatti, J. J. Beck, T. Patterson, R. Keil, J. D. Goldman, H. D. G., B. Baldwin, hitne,C 98 ntesseai oiinof position systematic the On 1928. C. Christensen, aie .M,S .Rsel .Sno-ura n .Francisco-Ortega. J. and Santos-Guerra, A. Russell, J. S. M., A. Carine, Macaronesian the of relationships Spatio-temporal 2006. M. A. Carine, of study taxonomic and morphological comparative A 1964. E. A. Bobrov, adi,B .adW .Wge.21.Hwia nisemradiations angiosperm Hawaiian 2010. Wagner. L. W. and G. B. Baldwin, arntn .S n .A ai.20.Rfgaadmgaini the in migration and Refugia 2007. Paris. A. C. and S. D. biogeogra- Barrington, fern in factors historical and Ecological 1993. S. D. Barrington, popu- avian on effects Evolutionary Phylogeographic 1998. 2001. Walker. D. Weigel. and C. D. J. Avise, P. and Browne, A. R. S., B. Arbogast, E.M.S.’s of The part from is University. S. article Duke M. at E. This Biology to in Taxonomists. Grant dissertation Plant Research doctoral Student of Research to Graduate of Society Grant-in-Aid a Grant Xi and American Improvement Sigma S., a Dissertation M. (DEB-1110775), E. S. Doctoral M. to E. NSF was and work a P. This M. by manuscript. K. this part of comments, in versions their Kao, earlier funded for T.-T. of reviewers Johnson, criticism anonymous A. and two Huiet, edits, and L. Rothfels, Russian Grusz, C. with A. Li, help to F.-W. for his grateful Manos collect for are P. to Gryganskyi We to A. S. translation. goes and M. gratitude musings E. Our biogeographic to study. his this 2010-7) in (number used Permit specimens Use several Free Products Forest a oe,H .adJ .Kdri.19.Teefc fQaenr climatic Quaternary of effect The 1998. Kadereit. W. J. and P. H. Comes, and geochronology, Tectonics, 1994. Dalrymple. B. G. and A. D. Clague, ye .S n .K rs.18.Lt icnia n ooeehis- Holocene and Wisconsinan Late 1987. Prest. K. V. and S. A. Dyke, Bayesian 2012. Rambaut. A. and Xie, D. Suchard, A. M. J., A. Drummond, eln,L .15.Pclaiiso h oubaRvrGreflora. Gorge River Columbia the of the Peculiarities of 1958. phylogeny E. L. and Detling, classification the On 1973. R. E. Sota, la De atn,G .adG asivc.19.Mtra neiac fthe of inheritance Maternal 1992. Yatskievych. G. and J. G. Gastony, phyloge- Molecular islands: oceanic on Evolution 2002. C. B. Emerson, eeCdsCroain 05 eunhrvrin48sqec analysis sequence 4.8 version Sequencher 2007. 2005. Klimas. Corporation. T. Codes S. Gene and Neale, Ramp M. J. Ranker, A. T. O., M. J. Geiger, li vlto ntefr genus fern the in poly- evolution interpreting ploid and delimitation species to approach diploids-first 2012. editors. California Wilken, H. D. as oaikForening Botanisk Dansk mrcnJunlo Botany of Journal in American continent and the Macaronesia of into floras: back-colonization colonizations Mediterranean multiple and for evidence Macaronesian molecular the of Relationships 2004. opportunity? of window or series 895–903. relictual 54: a flora: endemic Zhurnal of species the Botany fNrhAeia origin. American North of utrayhsoyo h e nln flora. England New the of history Quaternary phy. Sciences Biological B: Series process. London. speciation tree the and American lations North of biogeography Pleistocene squirrels. and genetics hne npatdsrbto n evolution. and distribution plant on changes II readings Press. Selected Hawai‘i Islands. of Hawaiian University Honolulu: the in of 5–40 Pp. history Chain. ral Volcanic Hawaiian-Emperor the of origin oyo h arnieIeSheet. Ice Laurentide the of tory 1.7. BEAST Evolution the and BEAUti with phylogenetics Madron Press. Academic in ferns 229–244 Pp. Polypodiaceae. 432–438. cnJunlo Botany of ferns. Journal cheilanthoid ican in genomes mitochondrial and chloroplast process. and pattern understanding Evolution to approaches netic 237–263. 41: otae n ro,Mcia,U .A http://www.genecodes.com. A. S. U. flora. Michigan, Arbor, Ann fern software. Hawaiian the of origins Brittonia and biogeography Molecular d.A .Jry .A rbe n .A hms London: Thomas. A. B. and Crabbe, A. J. Jermy, C. A. eds. , ora fBiogeography of Journal ˜ 5 223–234. 35: o 9 3–4.(ocwadLeningrad). and (Moscow 534–545. 49: 9 142–158. 59: 4 160–172. 14: d .Bree:Uiest fClfri Press. California of University Berkeley: 2. Ed. . 9 1969–1973. 29: 1 951–966. 11: ora fMammalogy of Journal Polypodium 9 716–722. 79: ieaueCited Literature 2 1–10. 22: 1 1070–1085. 91: .o h lr fteU.S.S.R. the of flora the of L. naso Botany of Annals 0 275–279. 20: h esnmna:Vsua lnsof plants Vascular manual: Jepson The 2 302–319. 82: h hlgn n lsiiaino the of classification and phylogeny The Astrolepis Ge 6:457–463. 265: rceig fteRylSceyof Society Royal the of Proceedings ´ gahepyiu tQuaternaire et physique ographie Convolvulus 0:849–879. 105: (Pteridaceae). rnsi ln Science Plant in Trends Polypodium Rhodora oeua ilg and Biology Molecular (Convolvulaceae). d .A Kay. A. E. ed. , 0:369–386. 109: Botanicheskii . die Af Udgivet Systematic Molecular natu- A Amer- Taxon 3: Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. afe,C . .E ots n .S ots 95.Pyoeyo the of Phylogeny 1995a. Soltis. S. P. and Soltis, E. D. H., C. Haufler, mecha- and Mode 2000. Therrien. P. J. and Hooper, A. E. H., C. the Haufler, and analyses Chromosomal 1991. Zhongren. W. and H. C. Haufler, 1995. Ranker. A. T. and H. C. Haufler, afe,C .adM .Wnhm 91 e pce fNrhAmeri- North of species New 1991. Windham. D. M. and H. C. Haufler, Matern 2000. Raquin. C. and J.-M. Guillon, afe,C . .D ida,adE .Rb.19b eiuaeevolu- Reticulate 1995b. Rabe. W. E. and Windham, D. M. H., C. Haufler, oidrjl,R,C .Hge,adC .Bie.21.Phylogenetic 2011. Bailey. D. C. and Hughes, E. C. R., Govindarajulu, 04 IE TA. OYOIMPYOEY1053 PHYLOGENY POLYPODIUM AL.: ET SIGEL 2011. Clark. H. D. and A. Gillespie, 2014] afe,C . .D ida,F .Ln,adS .Wimr.1993. Whitmore. A. S. and Lang, A. F. Windham, D. M. H., C. Haufler, wtui . .Ymzk,D .Bufr,adH ha 1995. Ohba. H. and Boufford, E. D. Yamazaki, T. K., Iwatsuki, Hulte u,L-. .W i .L ho,adC-.Wn.21.Frtisgt into insights First 2011. Wang. C.-N. and Chiou, W.-L. Li, F.-W. L.-Y., Kuo, divergence? promote glaciations Pleistocene the Did 2001. L. L. Knowles, eit .M 96 oegntccneune fieae,adterrole their and 1990. ages, ice Price. of consequences G. genetic Some 1996. M. M. G. and Hewitt, Kramer, U. K. Veldhoen, P. E., Hennipman, elne,D .18.Ntso ot mrcnferns. American North on Notes 1981. B. D. Lellinger, The 1971. A. F. Lang, of position systematic the On 1977. Ikebe. C. and S. Kurita, idbad .J,C .Hulr .P hrin n .Wles 02 A 2002. Walters. C. and Therrien, P. J. Haufler, H. C. J., T. the Hildebrand, in oscillations climatic of consequences Genetic 2004. phylogeography—or M. and G. zones Hewitt, hybrid Speciation, 2001. M. G. Hewitt, i .1997. from phylogeny J. estimating to Li, approach likelihood A 2001. O. P. Lewis, Kvac ag .A 99 e aefraseisof species a for name new A 1969. A. F. Lang, PartitionFinder: 2012. Guindon. S. and Ho, W. Y. S. Calcott, B. R., Lanfear, ilbad .1888. W. Hillebrand, ˇ ´ inst data. site tion vulgare Polypodium contrasting habitats. of tropical and Biology Implications temperate Species representing pteridophytes: ferns in in patterns speciation of nisms Botany of Journal allopolyploid American of origin relationships. ei nihsaogsse pce ftefr genus can fern the of species Journal Fern sister American among insights netic horsetail the diver- of speciation. patterns allopatric and gent diversity species cryptic reveal Mimosoideae) ini the in tion n ouaingntcaaye fdiploid of analyses genetic population and Look Closer A Chronology: and Extent Glaciations: nary USA Polypodium aclrplants vascular fern grasshoppers. montane in Ecology models refugial explicit of Tests Gymnospermae and Pteridophyta I. Vol Japan. ina oit fLondon of Society Linnean speciation. and divergence in in plants 203–230 Pp. Polypodiaceae. York New Committee. Editorial Press. America University North Oxford Oxford: of and Flora ed. 2, vol. Journal west. makinoi e hybrid new Sciences Biological B: Series Quaternary. time. and space in genes seeing oyoimpellucidum Polypodium data. character morphological discrete r ot America. North substitution ern and schemes 1695–1701. partitioning 29: analyses. phylogenetic of for models selection combined Republic). (Czech Bohemia northern of Repertorium Oligocene the from Journal of hss arne ass nvriyo Kansas. of University Kansas: Lawrence, thesis. hnrgm n aclrcryptogams vascular and phanerogams k .20.Anwfsi pce of species fossil new A 2001. Z. ek, ,E 1968. E. n, oyoimscouleri Polypodium Cystopteris matK p 4–6 in 447–462 Pp. . Madron o.1 d .Kbtk.Bri:Springer-Verlag. Berlin: Kubitzki. K. ed. 1, vol. 1 90–94. 71: 2 214–228. 92: 0 691–701. 10: Mxm)Fomin. (Maxim.) isseai nlsso h aainedmcfr species fern endemic Hawaiian the of analyses Biosystematic phylogeny. oyoimvulgare Polypodium p 5–5 in 356–357 Pp. . lr fAak n egbrn ertre:Amna fthe of manual A territories: neighboring and Alaska of Flora Polypodium ˜ qieu variegatum Equisetum 1:159–177. 112: hlspia rnatoso h oa oit fLondon. of Society Royal the of Transactions Philosophical o tnod tnodUiest Press. University Stanford Stanford: . ytmtcBotany Systematic mrcnFr Journal Fern American 5 223–236. 15: 1 235–254. 21: and lr fteHwia sad:Adsrpino their of description A Islands: Hawaiian the of Flora oyoimvulgare Polypodium ope:Isgt rmclrpatDArestric- DNA chloroplast from Insights complex: Madron eeomnsi utrayScience, Quaternary in Developments Polypodium oeua hlgntc n Evolution and Phylogenetics Molecular 5 361–374. 85: n t ypti congeners. sympatric its and rvdsisgt ocrigtesystematics the concerning insights provides 8 247–276. 58: Plpdaee n eae congenerics. related and (Polypodiaceae) ora fJpns Botany Japanese of Journal oyoimvirginianum Polypodium mrcnJunlo Botany of Journal American 8 624–629. 78: 4:183–195. 349: ˜ o 0 53–60. 20: lr fNrhAeianrho Mexico of north America North of Flora complex. ilgclJunlo h ina Society. Linnean the of Journal Biological lcain fteSer eaa California, Nevada, Sierra the of Glaciations ihcmet nterreticulate their on comments with , 0 110–119. 20: oeua Ecology Molecular (Schleich.). h aiisadgnr fvascular of genera and families The RbcL odn ilasadNorgate. and Williams London: . 1 7–23. 81: oeua ilg n Evolution and Biology Molecular lihrtneo hoolssin chloroplasts of inheritance al ope ntePcfcNorth- Pacific the in complex ytmtcBiology Systematic ytmtcBotany Systematic Polypodium eune rvd phyloge- provide sequences Polypodium oy:Kdnh Ltd. Kodansha Tokyo: . urn Genetics Current Leucaena t oi:Elsevier. Louis: St. . 0 537–549. 10: 2 39–49. 52: (Polypodiaceae). rmnorthwest- from (Polypodiaceae) 8 2049–2063. 98: (Leguminosae; o.15, vol. mrcnFern American mrcnFern American Pleurosoriopsis 0 913–925. 50: 9 556–566. 59: Polypodium 0 89–109. 20: 7 53–56. 37: Molecular lr of Flora Quater- h D. Ph. Feddes Plant . e,R 08 acd.y vial thttp://www.bioinformatics.org/ at Available in gapcode.py; 2008. 57–67 R. Pp. Ree, islands. California the of floristics The 1965. H. P. Raven, Haufler, H. C. Smith, R. A. Kennedy, from C. S. Available Geiger, O. v1.4. M. Tracer J. A., 2007. biota? T. Ranker, Drummond. Hawaiian J. the A. is and old A. How Rambaut, 2002. Clague. A. D. and P. J. 1994. Price, Short. J. M. and R. J. Press, 1991. E. C. Pielou, of sorus The 1974. Kott. S. L. and L. R. Peterson, 2003. D. Palmer, 1997. N. C. Page, a,D .18.Dvlpetadrgoa ifrnito fthe of differentiation Maldonado,J.E.,C.Vila regional and Development 1987. H. D. Mai, 2005. Maddison. P. W. and R. D. Maddison, atn .14.Plpod in Polyploidy 1947. I. Manton, uaVg,I,J .Tejero-Dı D. J. The I., 1964. Lang. Luna–Vega, A. F. and M. R. Lloyd, in numbers chromosome New 1963. M. R. Lloyd, biog 1753. C. and Linnaeus, Taxonomic 1981. L. S. and systems, Lindsay, breeding variation, Genetic 1999. Haufler. H. C. and J. Li, atn,P 93 e rae lnuexde glanduleux organes Les 1943. P. Martens, of Cytology 1951. I. Manton, atn,P 90 e aahssde paraphyses Les 1950. P. Martens, 1950. I. Manton, ikl .T n .R mt.2004. Smith. R. A. phylogenetic and T. for J. Testing Mickel, 1996. Kellogg. A. E. and J. R. Mason-Gamer, to .M,T aßn .P rir n .Shedr 09 e insights New 2009. Schneider. H. and Kreier, H.-P. Janßen, T. M., E. Otto, veg- current and historic estimated of comparison A 2005. T. Oberbauer, phy- Molecular 2007. Pryer. M. K. and Schneider, H. S., N. Nagalingum, 1995. C. R. Moran, CIPRES the Creating 2010. Schwartz. T. and Pfeiffer, W. A., M. Miller, America tion  Garden. Botanical Islands Barbara California Santa the Barbara: of Santa biology Philbrick. the N. R. on symposium the of Proceedings the Evolution of and evolution Phylogenetics and genus phylogenetics Hawaiian Molecular 2003. endemic Parris. S. B. and http://beast.bio.ed.ac.uk/Tracer. Sciences Biological B: Series London. of Society divergence. Royal recent suggest phylogeny and Geology Museum. History Press. Hawai‘i Press. University .Rvs 02 igorpia nlsso two of analysis Biogeographical 2012. Rivas. G. uoenvgtto uigteTertiary. the lution during vegetation European 4.08 v. evolution. character and eny London of Society Linnean Society. Linnean the of America. Journal Central and Mexico in (Polypodiaceae) complexes oeapcso h eeomn n tutr fprpye and paraphyses of structure and development sporangia. the of aspects some in nteont he ( shrew ornate the in sions ot America. North Journal Fern ( chickarees California Botany Hawaiian Systematic in diversification of patterns ge ( Press. University Cambridge Cambridge: serratum ofitaogmlclrdtst ntetieTiiee(Gramineae). Triticeae tribe Biology the Systematic in datasets molecular among conflict notepyoeyof Polypodiopsida). (Polypodiaceae, phylogeny the into Symposium Islands California Mexico. Sixth the Island, of Guadalupe ings of structure community etation hetero- the in genus evolution fern sporous morphological Moran, and C. relationships Garden. R. logenetic Botanical Davidse, Missouri G. Louis: Knapp, St. S. Riba. Sanchez, R. Sousa and M. eds. 1, vol. Workshop Environments Louisiana. Computing Orleans, in New Gateway (GCE). 1–8 the Pp. trees. of phylogenetic large Proceedings of inference for Gateway Science Press. Garden Botanical York New The .vulgare P. ´ rick/software.html. ographique. 3 190–201. 53: 6:79–91. 161: hcg:Uiest fCiaoPress. Chicago of University Chicago: . . ultnd aSocie la de Bulletin aainJunlo Botany of Journal Canadian var. 3 99–101. 53: aa‘’ en n enallies fern and ferns Hawai‘i’s h en fBianadIreland and Britain of ferns The rbeso yooyadeouini h Pteridophyta the in evolution and cytology of Problems pce Plantarum Species ultnd adnbtnqed l’E de botanique Jardin du Bulletin fe h c g:tertr flf ogaitdNorth glaciated to life of return the age: ice the After Polypodium rts enGazette Fern British virginianum 4 339–355. 24: 5 524–545. 45: ` n .K an.20.Tiatt eei subdivi- genetic Tripartite 2001. Wayne. K. R. and , Marsilea Tamiasciurus Polypodium Pleopelti ` oe ornatus Sorex ´ te oyoimvulgare Polypodium z .CnrrsMdn,M ed,and Heads, M. Contreras-Medina, R. ez, ` Adenophorus 6 337–347. 26: oaed oaiu eBelgique de Botanique de Royale . )I.Valeursyste .P.3935in 349–365 Pp. L. ytmtcBotany Systematic lr fMadeira of Flora h trdpye fMexico of Pteridophytes The oma:IpessLuetiSalvii. Laurentii Imprensis Holmiae: . oyoimvulgare Polypodium udrad iae Associates. Sinauer Sunderland: . n eae etoia genera Neotropical related and s ). oeua hlgntc n Evolu- and Phylogenetics Molecular ora fMammalogy of Journal :168–177. 9: nAmerica. in oyoimvulgare Polypodium 2 2283–2288. 52: orpi eainhp fBaja of relationships eographic ). oeua Ecology Molecular cld :Aayi fphylog- of Analysis 4: McClade (Grammitidaceae). Polypodium oouu nvriyof University Honolulu: . ln ytmtc n Evo- and Systematics Plant abig:Cambridge Cambridge: . :143–153. 6: oyoimvirginianum Polypodium . oyoimvirginianum Polypodium ´ ´ Polypodium odn h Natural The London: . aiu tre et matique Nature a Bruxelles a tat 2 16–25. 32: Nature lr Mesoamericana Flora Polypodium 6:2429–2435. 269: tl sous-espe la et (Polypodiaceae). rceig fthe of Proceedings 5:136. 159: 0:940–955. 106: 6:37. 167: 2 673–683. 62: e York: New . 0 127–147. 10: 2 225–262. 82: L. ope in complex ´ Molecular American 7 1–14. 17: Biological partition Proceed- species ,ed. ` ce : . Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. ots .E,A .Mri,J .MLcln .S ao,adP .Soltis. S. P. and Manos, S. P. McLachlan, S. J. Morris, B. A. E., 1997. D. Soltis. Soltis, S. P. and Strenge, D. D. Gitzendanner, A. M. E., D. Soltis, genus The 2005. Stech. M. and Fontinha, S. Esquivel, G. M. sequence M., in Sim-Sim, characters as on Gaps structure 2000. Ochoterena. crown H. and the P. of M. Simmons, Effects 2003. Bailey. G. M. and C. S. Sillett, ie,E . .D ida,A .Sih .J yr n .M Pryer. M. K. and Dyer, J. R. Smith, R. A. Windham, D. M. M., E. Sigel, hgn,L n .Hulr 2013. Haufler. of C. taxonomy and L. and Shugang, cytology the to Contributions 1961b. of G. taxonomy M. and Shivas, cytology the to Contributions 1961a. G. M. Shivas, Rodrı 04SSEAI OAY[oue39 [Volume BOTANY SYSTEMATIC 1980. H. R. Roberts, 1054 mt,A .adD Tejero-Dı D. and R. A. Smith, Schneider. H. and Ranker, A. T. Haufler, H. C. Kreier, H.-P. II. R., A. Baicalensi, Smith, Flora de Notulae 1974. V. Siplivinski, cutpl,E,P oal n .M re.20.Plastid 2006. Pryer. M. K. and Korall, P. E., Schuettpelz, radiation Cenozoic a for Evidence 2009. Pryer. from M. K. inferred and E. phylogeny Schuettpelz, Fern 2007. Pryer. M. K. and E. Schuettpelz, Haufler, H. C. Hildebrand, J. T. Cranfill, R. The Smith, R. 2006. A. Smith. H., Schneider, R. A. and Wilson, R. Kreier, H.-P. H., Schneider, cmkv .20.Anew A 2001. A. Schmakov, udl,R .adT .Pie 09 dpierdain nonadaptive radiation, Adaptive Scha 2009. Price. D. T. and J. R. Rundell, Taxonomic 2014. Carine. A. M. and Schneider, H. Robba, L. J., F. Rumsey, M. K. and Chiou, W.-L. Korall, P. Kuo, phylo- L.-Y. Bayesian Larsson, 3: A. MrBayes J., C. 2003. Rothfels, Huelsenbeck. P. J. and F. Ronquist, ¨ e,H 2005. H. fer, oia comparison. logical 06 oprtv hlgorpyo nlcae atr North eastern unglaciated of America. phylogeography Comparative 2006. the 353–373. America. from 206: North plants of of Northwest phylogeography Pacific intraspecific DNA Chloroplast novelties. nomenclatural and 43–58. 92: genus the of inition bio- and ecology, affinities. geographic relationships, Archipelago—molecular Madeira Plagiochila analyses. phylogenetic based forests. fern redwood epiphytic the of biomass Brittonia 04 eicvr of Rediscovery 2014. Press. Garden Botanical Missouri ora fteLnenSociety Linnean the of Journal of species Society Linnean the of Journal of species ferns. among relationships deep 897–906. for support improved Polypodium 2006. rastenii vysshikh China ffrsi nagopr–oiae canopy. USA angiosperm–dominated Sciences of an Academy National in ferns of genes. plastid three and 1037–1050. species leptosporangiate 400 tion plants. the epiphytic of of aspects diversification polygrammoid exploring of Grammitidaceae): phylogeny and the (Polypodiaceae Unraveling ferns 2004. Ranker. A. T. and southern in Polypodiidae) America. (Polypodiaceae, South ferns polygrammoid Synammia Publishers. nEooy&Evolution & speciation. Ecology in nonecological and speciation ecological radiation, reassessed. conundrum: continental a and uncertainty ferns. II eupolypod of radiation Biology internodes rapid atic short ancient and the rates, resolve fast to roots, deep Overcoming 2012. Pryer. models. mixed under Univer- inference genetic Cambridge Cambridge: Parnell. N. in A. Press. J. sity 280–303 and Pp. Waldern, systematics past. S. and the from ecology future of the change, for reconstruction lessons integrative forests: laurel an Tethyan of demise the and ´ guez-Sa 1 1041–1063. 31: Serpocaulon os –,e.Foao hn dtra omte.S.Louis: St. Committee. Editorial China of Flora ed. 2–3, vols. ´ ce,F n .Ary.21.Cnzi lmt changes climate Cenozoic 2011. Arroyo. J. and F. nchez, 6 278–286 66: oeua Ecology Molecular Polypodium Polypodium Dmr. uot Paiciaee eaiohtn)in Hepaticophytina) (Plagiochilaceae, Dumort. (Dumort.) . 1 490–509. 61: oaia ora fteLnenSociety Linnean the of Journal Botanical Taxon nga vdnefratmeaelnaeof lineage temperate a for Evidence enigma: lr fteAoe:Afedguide field A Azores: the of Flora oyoimmacaronesicum Polypodium 1 327–337. 11: ytmtcBotany Systematic mrcnJunlo Botany of Journal American 5 919–930. 55: Plpdaee,anwgnssgeae from segregated genus new a (Polypodiaceae), h enGazette Fern The oyoimcalirhiza Polypodium oaHedwigia Nova nErp n mrc:I.taxonomy. II. America: and Europe in 4 394–399. 24: nErp n mrc:I cytology. I. America: and Europe in Polypodium ´ z 2014. ez. 5 4261–4293. 15: 8 27–38. 58: 13–25. 58: ytmtcBiology Systematic oyoimscouleri Polypodium d.T .Hdisn .B Jones, B. M. Hodkinson, R. T. eds. , Polypodium 0:11200–11205. 106: 1 31–41. 31: Pleopeltis rmAltai. from oeua hlgntc n Evolu- and Phylogenetics Molecular 2 69–74. 12: 1 449–461. 81: ln ytmtc n Evolution and Systematics Plant Bioinformatics Plpdaee nMexico. in (Polypodiaceae) and oyoimmacaronesicum Polypodium p 3–3 in 838–839 Pp. . Plpdaee,aredef- a (Polypodiaceae), 0 255–261. 90: ekrhi:Margraf Weikersheim: . Turczaninowia .australe P. 9 369–381. 49: 7:449–460. 174: Plpdaee in (Polypodiaceae) oot Sistematiki Novosti atpA rceig fthe of Proceedings oaia Sciences Botanical 9 1572–1574. 19: aaprovide data morpho- a : Taxon Taxon Botanical Botanical :5–7. 7: lr of Flora System- Climate Trends 55: 56: tee .A n .Sofr 06 olsetbsdhptei testing hypothesis Coalescent-based 2006. Storfer. A. and A. C. Steele, ro,R .adA .Tyn 1982. Tryon. F. A. and M. R. Tryon, in 13–40 Pp. flora. Macaronesian the of Origins 1979. P. Sunding, enn .L n .A akr 03 urn ttso h en and ferns the of status Current 2013. Ranker. A. T. and L. A. Vernon, 1964. H. D. Valentine, British some in counts Chromosome Tejero-Dı 1963. Lang. parsimony F. and using C. analysis M. T. Phylogenetic Taylor, PAUP* 2002. L. D. Swofford, adrore,A,F .Rme,adM .Crn.20.Does 2007. Carine. A. M. and Rumsey, J. F. A., Vanderpoorten, anr .H 94 aahss Filicineae. Paraphyses: 1964. H. W. Wagner, 1998. Gibby. M. and Barrett, A. J. Rumsey, J. F. Russell, J. S. C., J. Vogel, ils .L n .J htae.20.Terfga debate. refugial The 2000. Whittaker. J. R. and L. K. Willis, vial.Nmnltrlatoiisaegvnfrtefrtseie of specimen first the GenBank for taxon. given (herbarium); each are authorities number order; Nomenclatural collection available. the and in collec- collector accessions (http://fernlab.biology.duke.edu): locality; number tion accession Database 0.951. vers. GARLI, 2006. J. D. Zwickl, model a phylogeography: Amphibian 2008. Beebee. C. J. T. and I. Zeisset, substi- nucleotide of Rates 1987. Sharp. M. P. and Li, hybrid W.-H. novel H., K. A Wolfe, 2005. Yatskievych. G. and D. M. Windham, tetraploid the of Recognition 1991. Smith. R. A. and A. radia- rapid S. ancient Whitmore, Deciphering 2007. Lockhart. J. P. and B. J. insects: Whitfield, of radiations rapid Ancient 2008. Kjer. M. K. and B. J. Whitfield, Pleisto- 2009. Bond. E. J. and Marek, E. Scale. P. Stockman, Time K. Geologic A. GSA M., 2009 J. 2009. Walker, Geissman. W. J. and D. J. Walker, F6821 ,- -. -, -, EF463832.1, le oaice atnGoettingen:; Garten Botanischer Alter irsrmvarians Microsorum rmteimmiyoshianum Drymotaenium ule Co.: Hualien atuCo.: Nantou -. GQ256255.1, -, -, Appendix rmteimmiyoshianum Drymotaenium uprsmlil litcn eui ntePcfcNrhetfor Northwest Pacific ( the salamander giant in Pacific refugia the Pleistocene multiple supports colpodes eeec otoia America tropical to reference 75–115. 67: islands and ogy redefinicio ferns. Columbia Associates. Sinauer Sunderland: 10. beta 4.0 v. methods), other (*and aaoei xs?Cnlcigsga ntebypyeadpterido- and bryophyte the floras. in H. phyte signal V. A. Conflicting D. exist? Edmonson, Macaronesia and R. Walters, J. M. Charter, S. Press. Valentine, O. University H. Cambridge A. Cambridge: D. Webb. Burges, Moore, M. A. D. N. Heywood, Tutin, G. T. vdnefrmtra rnmsino hools N nthe in DNA chloroplast of 247–249. transmission maternal genus for Evidence Islands. Hawaiian the 59–112. of lycophytes h hlgntcaayi flrebooia eunedtst under datasets sequence Texas. criterion. biological of likelihood large maximum of the analysis phylogenetic the 109–119. distributions. 101: species of aspects historical understanding for and chloroplast, mitochondrial, 9054–9058. 84: plant DNAs. among nuclear greatly vary tution Arizona. from (Polypodiaceae) 1406–1407. Madron calirhiza Polypodium tions. 449–472. analysis. 53: phylogenetic for challenges Biology data. genus paleoclimatic and coastal millipede phylogeographic, and genetic, the Mountains Appalachian in the plain across refugia glacial cene Today GSA ´ 5 2477–2487. 15: z .D n .Pceo 04 aaneo,nomenclatura, nuevos, Taxa 2004. Pacheco. L. and D. J. ez, rnsi clg Evolution & Ecology in Trends ˜ slnu ( Asplenium o :25–36. 9: uz Plpdaee Pteridophyta). (Polypodiaceae, Kunze .Seiesicue nti td.Txnnm Fern name Taxon study. this in included Specimens 1. 8 233–248. 38: .Shetez1074A Schuettpelz E. ´ .Shetez1136A Schuettpelz E. d .Bawl.Lno:Aaei Press. Academic London: Bramwell. D. ed. , distribucio y n 9 60–61. 19: mrcnJunlo Botany of Journal American irsrmfortune Microsorum rceig fteNtoa cdm fSine USA Sciences of Academy National the of Proceedings mrcnFr Journal Fern American Mt. enpa Hett. & Hennipman (Mett.) heoimdecumanum Phlebodium Polypodium slnaee Pteridophyta). Aspleniaceae, Plpdaee nwsenNrhAmerica. North western in (Polypodiaceae) tA aK rbcL, matK, atpA, ´ elsepce eaindscon relacionadas especies las de n - :CHINA.Sichuan: e ok Springer. York: New . p 51 in 15–16 Pp. . iapoo tenebrosus Dicamptodon Mkn)Makino (Makino) mrcnFr Journal Fern American Narceus (DUKE); DK) F008 F003 ,-. -, KF909023, KF909068, (DUKE); en n lidpat ihspecial with plants allied and Ferns hD hss utn University Austin: thesis. Ph.D. TMoe Ching (T.Moore) 2 258–265. 22: eei loih prahsfor approaches algorithm Genetic 3 123–126. 53: .Shedrs n s. Schneider H. nulRve fEntomology of Review Annual vdnefo population from Evidence : 4 625–638. 94: Taxon mrcnFr Journal Fern American and -, lr Europaea Flora F004 F004 -. KF909054, KF909024, .C i DB06104 Liu C. C. Wld)J Sm. J. (Willd.) 3 56–64. 13: trnG-R caBtnc Mexicana Botanica Acta 3475: oaiaActa Botanica M Evolutionary BMC 4879: ). 4817: aanot data = - . 5 57–67. 95: oeua Ecol- Molecular ncultivation, In Science Polypodium Polypodium (GOET); . TAIWAN. TAIWAN. o.,eds. vol.1, Heredity Plants 2384: (PE); 103: 111: 287: Delivered by Publishing Technology to: Duke University IP: 152.3.102.242 on: Fri, 17 Oct 2014 14:51:56 Copyright (c) American Society for Plant Taxonomists. All rights reserved. F007 ,- -. -, -, KF909087, n. plesiosorum s. Vernon L. A. martensii 47 Larsson macaronesicum Polypodium Columbia: KF909111. KF909051, glycyrrhiza BG04-129 Barker F003 KF909113. KF909053, KF909109. Washington:KF909049, A. S. U. KF909108. KF909048, -, KF909078, (DUKE); glycyrrhiza 2010-89 Polypodium Sigel M. E. 9399 F005 F626 -. EF463256, KF909025, oyoimglycyrrhiza Polypodium Fukui: 1055 KF909101. KF909040, -, KF909083, PHYLOGENY POLYPODIUM AL.: ET SIGEL Napo: ECUADOR. 2014] F007 ,K994,KF909104. KF909044, -, KF909067, -: Goettingen: Garten Botanischer -. oyoimcolpodes 8-61 Polypodium Number: lection cambricum Polypodium ENGLAND. KF909115. KF909056, KF909027, makinoi KF909086, (DUKE); J2661 -. FJ825696.1, oyoimamorphum Polypodium 2010-104 KF909095. Washington: KF909094. KF909034, KF909009, Windham & Andrews E.G. AAA rts Columbia: British CANADA. KF909116. KF909057, KF909028, EIO eaCruz: Vera MEXICO. 926 Haufler H. C. rts Columbia: British -. -, KF909007, KF909069, (DUKE); GE) F680 F006 F628 -. EF463258, KF909026, EF463840, Mexico:(GOET); from originally Goettingen, Garten Botanischer sanctae-rosea Polypodium -. KF909100. KF909038, -, -, (DUKE); appalachianum 38496Polypodium Oldham J. M. KF909098. Virginia: KF909097. Carolina: KF909035, North A. S. U. -. KF909031, AAA Ontario: CANADA. 98-109 Caswell KF909119. KF909063, Grev .S .California: A. S. U. re 20)ada eitdi i.3 g osrit oenme in number node constraint: Age 3. Fig. A in and depicted 3 Fig. as and (2009) Pryer Pahang: 12-05 KF909121. -. Hidalgo: KF909060, MEXICO. -, KF909092, KF909120. sibiricum ue 141 Huiet KF909117. KF909058, -, .B es .n. s. Sessa B. E. oyoimcalifornicum Polypodium Appendix ltcru stemaria Platycerium U) ,- F006 -. KF909046, -, -, (UC); 7216: TI) ,- Y6351 -. AY362345.1, -, -, (TAIF); .Eiaa2973 Ebihara A. .Shetez786 Schuettpelz E. Mkm)Fomin (Makim.) DK) F000 F009 F009 KF909118. KF909059, KF909029, KF909090, (DUKE); Sipliv. .J ohes3905 Rothfels J. C. DK) F004 F002 F003 KF909112. KF909043, KF909022, KF909084, (DUKE); Mett. oyoimsibiricum Polypodium rspi contigua Prosaptia .S .Washington: A. S. U. .J ohes4046 Rothfels J. C. oyoimamorphum Polypodium oyoimappalachianum Polypodium DK) F003 F000 F003 KF909096. KF909033, KF909010, KF909073, (DUKE); 8161: PPENDIX .S cafes n s. McHaffie S. H. Kunze oyoimrhodopleuron Polypodium .Nd g osrit bandfo cutpl and Schuettpelz from obtained constraints age Node 2. oyoimappalachianum Polypodium .M ie 2010-125 Sigel M. E. AA;- ,K996,-. KF909061, -, -, (ALA); WS;- ,- JN189024. -, -, -, (WIS); 7540: 8008: KN) ,- 2151 -. U21145.1, -, -, (KANU); .plesiosorum P. AA;- F009 F000 KF909110. KF909050, KF909019, -, (ALA); DK) F005 ,K995,KF909114. KF909055, -, KF909085, (DUKE); .S .Alaska: A. S. U. .Mtgr176 Metzgar J. .Shetez216 Schuettpelz E. .J ohes4065 Rothfels J. C. ;crepnignd ubrfo cutpl and Schuettpelz from number node corresponding 3; .J ohes3026 Rothfels J. C. DK) F000 F001 F007 KF909107. KF909047, KF909021, KF909080, (DUKE); DK) F006 F003 F007 KF909099. KF909037, KF909013, KF909076, (DUKE); .R rne 1708 Brinker R. S. oyoimpellucidum Polypodium oyoimglycyrrhiza Polypodium 7160: Kunze .S .Alaska: A. S. U. EIO Quere MEXICO. oyoimglycyrrhiza Polypodium oyoimsibiricum Polypodium .M ie 2010-85 Sigel M. E. DK) F007 F006 F005 KF909106. KF909045, KF909016, KF909077, (DUKE); E;K996,K991,K994,KF909103. KF909042, KF909015, KF909066, (E); .Hit145 Huiet L. ltcru superbum Platycerium oyoimscouleri Polypodium oyoimappalachianum Polypodium .M ie 2010-6 Sigel M. E. 8210: 7781: 8787: ..Eaton D.C. 8723 DK) F682 ,- -. -, -, EF463842, (DUKE); oyoimglycyrrhiza Polypodium Bav)Desv. (Beauv.) EIO Jalisco: MEXICO. 7249: Mxn .Chr. C. (Maxon) 7254: 8045: A.E.Bobrov DK) F005 F002 KF909036, KF909012, KF909075, (DUKE); 6489: DK) F001 F007 F002 -. KF909052, KF909017, KF909081, (DUKE); oyoimsubpetiolatum Polypodium AA.Fukui: JAPAN. : .Presl C. .S .Washington: A. S. U. .D ida 3404 Windham D. M. .S .Washington: A. S. U. PI.Valencia: SPAIN. .D ida 94-115 Windham D. M. E;K996,- F001 KF909102. KF909041, -, KF909065, (E); . -: DK) F002 F004 KF909039, KF909014, KF909082, (DUKE); 9145: oyoimamorphum Polypodium .S .California: A. S. U. oyoimfauriei Polypodium oyoimamorphum Polypodium EIO Chiapas: MEXICO. rspi contigua Prosaptia oyoimamorphum Polypodium .S .NwYork: New A. S. U. .P re .n. s. Krier H.-P. MEXICO. oyoimcalifornicum Polypodium EIO Jalisco: MEXICO. .Lpi 04-286 Lipkin R. 7773: DK) F008 ,- KF909105. -, -, KF909088, (DUKE); 7537: .L akr9347 Parker L. C. DK) F6861 ,EF463256.1, -, EF463836.1, (DUKE); ´ oyoimcambricum Polypodium DK) F002 ,KF909032, -, KF909072, (DUKE); JAPAN. taro: loetspolypodioides Pleopeltis 293: DK) F003 ,KF909064, -, KF909093, (DUKE); DK) ,K993,KF909062, KF909030, -, (DUKE); Kunze 8688: DK) F000 KF909008, KF909070, (DUKE); 8029: oyoimrhodopleuron Polypodium DK) F004 KF909011, KF909074, (DUKE); 1 DK) F009 KF909018, KF909079, (DUKE); .S .Oregon: A. S. U. oyoimscouleri Polypodium oyoimglycyrrhiza Polypodium .S .California: A. S. U. oyoimsibiricum Polypodium TAIWAN. .Reosi53471 Rzedowski J. 3544: afe Windham & Haufler .Lunr01-39 Lautner J. Kaulf. 7251: 8009: 8010: 3580: PI.Cnr Islands: Canary SPAIN. eJnh Hennipman & Jonch. de .B ek1160 Beck B. J. .Eiaa2972 Ebihara A. .H afe .n. s. Haufler H. C. 610: AAA oaScotia: Nova CANADA. 8523: arc ae lt Col- Blyth James Patrick .S .California: A. S. U. ncliain Alter cultivation, In .S .Alaska: A. S. U. 7768: 8778: EIO eaCruz: Vera MEXICO. .Hit138 Huiet L. ncliain Alter cultivation, In .S .Alaska: A. S. U. .Hit144 Huiet L. AA;- KF909020, -, (ALA); .M ie 2010-101 Sigel M. E. Christ DK) KF909071, (DUKE); GE) EF463837, (GOET); .M ie 2010-75 Sigel M. E. AA;K999,-, KF909091, (ALA); DK) KF909089, (DUKE); .J lhm27283 Oldham J. M. 4204: AAA British CANADA. .J ohes3031 Rothfels J. C. .L ho 97-09- Chiou L. W. .Shetez525 Schuettpelz E. 7630: 8499: .S .Oregon: A. S. U. Pleurosoriopsis .S .Hawaii: A. S. U. 7771: 8722 Sudsk. Hook. MALAYSIA. GE) ,-, -, (GOET); Polypodium Polypodium Polypodium Polypodium Kaulf. .M Sigel M. E. CANADA. DK) -, (DUKE); U) ,-, -, (UC); .S A. S. U. JAPAN. : .Toren D. L. ok & Hook. (Weath.) (DUKE); (DUKE); (DUKE); .S A. S. U. (DUKE); .H. M. 7533: 6951: 8786: 8039: 3829: 6485: 7783: 7255: A. P. L. -: .0mya. 3.00 5.751812) 42631(51.9024427-144.3365683). 94.2568391 (56.7785-148.1528); HPD); recovered. (95% not age Node = node NA mean HPD). 3 (95% age Analysis node HPD); mean (95% 4 Analysis HPD); Analysis age (95% node age node mean mean 1 2 Analysis number: Node 3. Figure .23(.751.33;581 289-.94;87525(4.8365912- 8.7155285 (5.4489-13.4990); (2.8599-9.2974); 5.8812 9.2201 12.8247527). (4.7765-13.0323); (5.0561-13.0767); 8.7273 (5.1362716-12.8036577). 8.8067 8.8091156 (6.845854-16.1018793). (5.3521-13.4785); 11.3841674 (7.7775471-16.2409955). (5.5573-13.4450); 11.8900939 13 (8.5236-17.1183); Node 12.6647 12.6441798 16.7618); (9.2312-18.0843); (10.3315- 13.4673 (8.489163-17.1542618). (8.1951-17.6302); 14.4694 12.6786 (8.9830-18.6721); (9.8967262-10.098109). 18.4595); 13.6208218 13.5960 19.2985); (9.9477-19.3999); (11.0786795-17.97002). 14.5497633 14.4076 (14.6166243- (11.2866-18.1274); 20.4979249 14.6256 (16.7382-27.2800); 26.8414717). 22.0444 (14.0057-27.0553); 24.6547965 (25.5001-31.2645); (17.1705195-32.3178474). 27.6381 (25.8946- (16.7244-32.7388); 24.7731 29.7374 31.4888); (25.856626-33.4618959). (25.8864-33.5223); 29.6668509 29.6928 33.4966); (32.8910391-39.154408). (25.9940-33.5855); 43.284612 36.0076033 29.7660 (36.0497452- (32.9915-39.2393); 39.0308866 36.1174 (39.5048-47.0098); (36.3638-42.0730); 41.9501179). 39.2372 43.3403 (36.1520-41.9606); (39.3988-46.8713); (39.4220265-46.9229102). (48.2060-77.2235); 43.2360 61.5822 (47.6548397-77.0995208). (46.6538-71.3184); 60.8147904 58.1556 (47.5398-73.5632); re 20,Fg 1;bs g siaefo cutpl n re (2009) Pryer and Schuettpelz deviation. from standard estimate one age ± best S1); Fig. (2009, Pryer mya. .19(.53645) .719 (2.5218709-9.076938). 5.5721493 (1.7563-6.4851); 4.0109 .22;266 105-.11;26397(1.2691906-4.2842092). 2.6533937 (1.0553-4.4121); 2.6264 4.1212); 3.8198199 (1.3417-4.4725); 2.7677 (1.6698643-6.4611226). (1.5281-6.2363); 3.7170 (1.5731-6.5035); .249 (0.4548203-1.8930163). (0.519568-1.9518167). 1.1294496 1.1977087 (7.7165-16.9081); 26 1.2080 1.2503524 1.8584); (0.2998-2.3730); 1.3046 (0.2954099-2.3214561). (0.2884-2.1737); 1.1538 (0.3471-2.2985); (0.3823619-2.3047871). 1.2580641 (0.4292-2.3980); 1.3331 (0.6166255-2.3069672). 1.3994 (0.9464934-4.084822). 2.3351); 2.3593704 21 (1.0302-4.2018); (0.6213-2.4462); Node 2.4402 1.4515 (0.8579-3.8495); (0.9421-4.1207); (0.9935786-4.2383673). 2.3787 2.491035 (0.9812-4.2770); 2.5202 000-.43;017 000-.15;013 000-.73;0.1590446 (0.0000-0.4753); 0.1634 (0.0000142-0.4614606). (0.0000212-0.6317393). (0.0000-0.4155); 0.1473 0.2150414 (0.0000-0.4443); -. -; -; (0.0001-0.6561); (0.0000075-1.0251555). (0.0000-0.8115); 0.3333279 0.2737 (0.0002-0.7849); -. 0.3028 -; (0.0102414-1.0037977). 0.9362); 0.4373267 (0.1611844-1.2026506). -; (0.0987683-0.9673032). 0.8916); 0.4863264 0.6384346 1.0147); (0.1325-1.3343); (0.0971099-1.2048526). 34 0.6801 Node 1.0108); (0.044884-1.5962671). 32 Node (0.1580288-1.3681133). 31 0.7077625 (0.2437-1.4460); Node (0.0470-1.6873); 0.7821 0.7303 (0.0428-1.4611); (0.0498-1.5583); (0.122964-1.4402922). 0.6668 0.7665087 (0.0406-1.6038); (0.202273-1.468254). 0.7808707 0.6885 (0.2796-1.5034); 0.8214 -. 1.3963); -; (0.2565-1.3793); .61(.35210) .56(.90226) .877 (0.5851661- 1.3897871 (0.5990-2.2961); 1.3546 2.3502674). (0.5335-2.1206); 1.2661 .07(.65183) .39(.33173) .74(0.4971-1.9486); 1.1774 (0.4333-1.7530); 1.0339 (0.4685-1.8332); 1.1057 : oe1 Node Appendix A: C: oe3 oe37 30 .0mya. 4.30 ± 43.00 357; node 3; node oe5 oe39 70 .0mya. 3.70 ± 37.00 359; node 5; node .73(.01237) .92(.12215) .33(0.6289- 1.4353 (0.5162-2.1958); 1.2912 (0.6031-2.3371); 1.3733 : ;054 014-.29;- .087 (0.0393405-1.6279652). 0.7018873 -; (0.1547-1.1289); 0.5845 -; : .83(.06115) .47(.94092) ;0.5976591 -; (0.0904-0.9022); 0.4497 (0.1066-1.1952); 0.5883 0.6913805 : -; (0.1052-1.1135); 0.5465 (0.1576-1.2059); 0.6314 : 227 754-744) 133 684-654) 9.3523 (6.8946-16.5744); 11.3832 (7.5440-17.4243); 12.2370 : F: 099 4.631671) 858 4.271330) 99.3296 (48.3207-113.3601); 78.5680 (49.8643-116.7018); 80.9093 : oe9 oe31 42 .2mya. 1.42 ± 14.20 361; node 9; node oe9 Node 5 Node oe23 Node oe16 Node .Dvrec ieetmtsi Y o oe in nodes for MYA in estimates time Divergence 3. oe38 Node 467 1.121.54;1.00(11.1799-18.0744); 14.5090 (11.2182-18.0584); 14.6571 : (32.8590-39.1146); 36.0304 (33.0286-39.2506); 36.0998 : oe28 Node oe33 Node .62(.16222) .90(0.3919-2.2016); 1.1960 (0.4176-2.2920); 1.2622 : oe12 Node oe35 Node .49(.72910) .93(2.4591-8.9299); 5.4933 (2.4742-9.1701); 5.7479 : oe18 Node oe25 Node .60(.05061) .05(0.0000-0.9507); 0.3085 (0.0065-0.6118); 0.2690 : oe8 Node 4 Node oe40 Node 267 809-715) 199 (7.4503- 11.9199 (8.0990-17.1256); 12.6070 : .34(.47134) .56(0.1806- 0.7586 (0.1427-1.3746); 0.7304 : .10(.70137) .15(0.1347- 0.5105 (0.0770-1.3073); 0.6120 : ;042 003-.52;051 (0.1174- 0.5012 (0.0131-0.9512); 0.4024 -; : .58(.82439) .19(1.1450- 2.5169 (1.2852-4.3195); 2.6588 : .58(.93181) .12(0.4810- 1.1152 (0.4983-1.8718); 1.1598 : oe20 Node oe30 Node oe27 Node 227 1.672.55;20.5928 (16.5627-27.5515); 22.2375 : 39.0101 (36.2754-41.9505); 39.1587 : oe15 Node oe3 Node oe22 Node .31(.00068) 0.2202 (0.0000-0.6688); 0.2301 : B: .95(.54401) 2.1815 (0.9574-4.0619); 2.2995 : .88(.30153) 0.6425 (0.0350-1.5430); 0.6818 : .82(.31163) 0.7914 (0.4331-1.6731); 0.9822 : oe36: Node oe4 oe38 92 3.92 ± 39.20 358; node 4; node D: oe37 Node 323 (39.5811-47.0155); 43.2737 : oe11 Node oe7 Node .08(5.2464-13.8106); 9.3008 : oe6 oe32 00 ± 30.00 352; node 6; node .34(0.5642-2.1744); 1.3384 : ;032 (0.0000- 0.3428 -; : 782 (25.6005- 27.8021 : oe14 Node 341 (9.0761- 13.4419 : ;039 (0.0165- 0.3992 -; oe41 Node oe17 Node oe24 Node oe2 Node oe39 Node oe10 Node oe29 Node oe19 Node oe6 Node 59.0926 : 9.3332 : 0.1551 : 3.9132 : 1.2200 : Node :-; : : : :