Phylogeny of Haplolepideous Mosses

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Published by Maney Publishing (c) British Bryological Society O 10.1179/1743282012Y.0000000014 DOI ß ottemnpyyo irndeadaposition a and Dicranidae of sup- monophyly far All the mouth. so port capsule reconstructions the phylogenetic around teeth molecular of row single a oaMRos M Rosa auai,ScinNN ednUiest,P o 54 30R Leiden, RA [email protected] Biodiversity 2300 Email: 9514, for Box Netherlands. PO Centre The University, Leiden Netherlands NHN, Section Stech, Naturalis, Michael to: Correspondence are arthrodontous–haplolepideous They ( the ecosystems. by terrestrial all are characterized almost and Dicranidae in diverse occur highly (Bryophyta). ecologically largest and mosses second morphologically of the or form subclass mosses haplolepideous (Dicranidae) the haplolepids 2009), families Stech, 30 and & genera (Frey 232 in species 4000 about With Introduction Keywords: 1 Stech Michael perspectives and — challenges mosses haplolepideous of Phylogeny o h ifrn ye fhpoeieu eitms utemr,gntcadgnmcrsac using the research of genomic and Implications genetic understudied. Furthermore, reviewed. peristomes. discussed severely briefly to haplolepideous exemplarily is also are of taxa are but haplolepideous types evolution that phylogeny, different character at genera Dicranidae the morphological necessary the for for complex are of reconstructions the taxonomically markers backbone phylogenetic of of the molecular and combinations resolve part large new to in on especially tackle support based levels, statistical analyses taxonomic significant phylogenetic different receive further as clades that well as of concluded respective Dicranales position of the families The Pottiales; most analyses. namely speciose mosses, haplolepideous most the the of Grimmiales, portion includes largest clade, the unsupported comprises and or grade Eustichiaceae, a Archidiaceae, as appears Leucobryaceae, which group, second new The reconstructions. a with as well as ( genomes are markers are different plastid and markers non-coding from of chloroplast markers combination mosses few novel on on a of based based on reconstructions based approaches group phylogeny phylogenetic mosses. recent haplolepideous molecular largest the with early addresses in of compared and relationships second systematic state results unravelling current main in the the The problems summarizes remaining review form urgent This most diverse. the (Dicranidae) highly ecologically and mosses morphologically haplolepideous The Germany eBiologı de ilg,Uiest fFoia ansil,F,USA, FL, Gainesville, Florida, of University Biology, Tecnolo ihnDcaia.Tefrtgopcmrssmrhlgclydvreseisfo ifrn families different from species distinguished diverse provisionally morphologically are comprises groups Ditrichaceae major group Distichiaceae, three first Catoscopiaceae, data, (Bryoxiphiaceae, The molecular available Dicranidae. the within to According spacer). Rhabdoweisiaceae ehrad etefrBoiest auai,ScinNN ednUiest,TeNetherlands, The University, Leiden NHN, Section Naturalis, Biodiversity for Centre Netherlands Dicranum rts rooia oit 2012 Society Bryological British ´ iaIndoame gica irnde allpdosmse,Nncdn lsi akr,Pyoey Review Phylogeny, markers, plastid Non-coding mosses, Haplolepideous Dicranidae, ´ eea (Bota Vegetal a tp)prsoe hc sal osssof consists usually which peristome, -type) 4 lfWerner Olaf , 1 ´ p.p. tatFMcDaniel F Stuart , ia mao Ecuador, Ambato, rica, n Scouleriaceae and , ´ ia,Uiesddd uca apsd siad,Spain, Espinardo, de Campus Murcia, de Universidad nica), Amphidium 4 Jesu , nbtentescn n hr ru ean miuu.I is It ambiguous. remains group third and second the between in ´ 6 p.p. esIsiuefrBoiest fPat,Uiest fBonn, of University Plants, of Biodiversity for Nees-Institute Mun s 2 aalHerna Rafael , ,wihfr rdsbacigoffrti h phylogenetic the in first off branching grades form which ), 3 elJardı Real ˜ alnaglaucescens Saelania oz ubro allpdostx naaye fa (Goffinet of mosses analyses of in range taxa haplolepideous broader of number Hedderson aFarge 1999a,b; fo- (Stech, La explicitly mosses haplolepideous either the studies on cussed These inferred. Dicranidae be the could ordinal within which relationships family-level from and published lineages, moss were major analyses the several of analyses phylogenetic of ancestor. diplolepideous peristome a haplolepideous from evolved the that indicate therefore Cox and 2008 Frey, the & arthrodontous–diplolepideous mosses, Stech in of (reviewed subclass Bryidae largest diplolepideous-alternate the the to sister as mosses, within nested 3,5 ntefis afo h atdcd,aboigperiod booming a decade, last the of half first the In ´ Bota n ita Quandt Dietmar , ´ p.p. io(SC,Mdi,Spain, Madrid, (CSIC), nico tal. et ´ tal. et ndez-Maqueda rmodaee Pottiaceae Drummondiaceae, , ora fBryology of Journal 00 02 Tsubota 2002; 2000, , 04 ricue considerable a included or 2004) , rps Dtihca) h hr group third The (Ditrichaceae). tal. et 4 -trn 00.Temlclrdata molecular The 2010). , einand region F 6 5 Universidad 2012 3 , 2 4 eatetof Department Departamento tal. et tal. et VOL atp .34 ,2001; B 2003; , -rbc p.p. NO .3 L , 173 Published by Maney Publishing (c) British Bryological Society tc tal. et Stech 174 ora fBryology of Journal tnadmresi ospyoeeis( phylogenetics moss in markers standard rbc eune ntoparts, two in sequenced The DNeasy extracted Germany). was the DNA genomic with using Total cleaned thoroughly water. were distilled shoots of parts sequencing Distal and PCR, extraction, DNA as well of species as 54 comprised Dicranidae sampling taxon data present sequence The of compilation and material Plant Methods and Materials ( regions DNA plastid three of more Tsubota hlgn fmse yCox by mosses of phylogeny loci mitochondrial novel (Wahrmund on resolving based approaches to recent much However, contribute Dicranidae. within not relationships higher-level did which (e.g. 2008), published were general in Qiu plants land or mosses of phylogenies new few only Thereafter 2010). Quandt, ae ihanvlpyoeei eosrcino a of reconstruction phylogenetic 50 novel com- a are with approaches pared back- earlier ambiguous be The remaining relationships. should the bone strategy resolve to which followed particular future in discusses (supra-) and challenges, Dicranidae to within ordinal levels at generic relationships phylogenetic on tackling phylogeny. of Dicranidae period of new problems a remaining of the beginning the mark could genomes three all ( from markers on based Goffinet both by (2011), Dicranidae in inference phylogenetic osbe isn eune of sequences missing possible, studies, mentioned available to addition In trn (Appendix). used were L collections herbarium from additional as well as (2008) Frey Stech 2004), (1999a, Stech from Specimens representatives. outgroup as lyptidae) and (Timmiidae) osmdltx sbifl reviewed. briefly is taxa haplolepid- model using eous research the genomic and for genetic more, Further- discussed peristomes. haplolepideous of exemplarily types different are cha- morphological evolution for racter data molecular available the al. et the olw ry&Seh(2009). Stech & Cla Frey follows (Appendix). from sequences some with GenBank completed In was study. dataset present the the cases, for generated newly were rps z hspprrvestecretsaeo knowledge of state current the reviews paper This - and L-F )ta r tl mn h otcmol used commonly most the among still are that L) hlgn fhpoeieu mosses haplolepideous of Phylogeny ,mitochondrial 4, rps tal. et 08)and 2008b) , ao e ae nnncdn lsi markers, plastid non-coding on based set taxon 4 -trn tal. et 06 tc unt 06 tc Frey, & Stech 2006; Quandt, & Stech 2006; , T tal. et -trn atp 04.Te eealbsdo n or one on based all were They 2004). , B L rps nayt streptocarpa Encalypta 09 00 swl stegenus-level the as well as 2010) 2009, , -rbc -trn 2012 atp rps 4 H nad -trn eune rmteabove the from sequences L ein(Herna region F 4 B -trn rps ln i Qae,Hilden, (Qiagen, Kit Plant ,adncerrbsml26S), ribosomal nuclear and 5, -rbc einwsapie and amplified was region F VOL 4 eune n,a a as far as and, sequences L imaaustriaca Timmia -trn tal. et pcr mlctosof Implications spacer. L sfiaino Bryophyta of ssification .34 and L trn 20)adSeh& Stech and (2006) - and L-F tal. et NO trn .3 trn ńdez-Maqueda L-F, ew (Enca- Hedw. -.Tefirst The L-F. 21)and (2010) cf. atp rps tc & Stech B Hedw. ,and 4, -rbc tal. et L eearayaalbefrprso h ein(e.g. region the of parts for trn numbers available accession already earlier were if specimen; per accession l eune sdi hssuyaegvni the in given are study this in The Appendix. used for sequences numbers accession all GenBank PCR. were for primers used Sequencing those well. as performed was sequencing automated Inc. the Macrogen where by (www.macrogen.com), or USA) WI, Madison, Wizard (Promega, the using purified were trn 3 the comprised part mlfidadsqecdsprtl ihprimers with respectively (Herna separately A-Fbryo/P6/7, and sequenced rps4-166F/A-Rbryo and amplified the products, PCR rps obtaining with difficulties of cases N eune eemnal lge nPhyDE in aligned (Mu manually v0.995 were sequences DNA phylogenetic reconstructions and analysis, sequence Alignment, pne h complete the spanned rmr tB1adrc- yChiang by rbcL-1 and atpB-1 primers yTaberlet by trn p416 n P6/7), and rps4-166F studies: previous rps in described as were used primers and oteemdlbsdaaye,mdltsigwas testing model analyses, model-based these Prior 2001). to using Ronquist, method, & (Huelsenbeck Metropolis- Carlo v3.1 the MrBayes Monte on chain based Markov calculated coupled were probabil- (PP) posterior ities Bayesian PAUP. with performed effect. in options same pseudo- the bootstrap with per replicate cycles addition random 10 replicates and 1000 with 10 performed were chosen superimposed searches parsimony under bootstrap and randomly Heuristic cycles. 2) addition of random to 25% up-weighted default iterations, positions the (200 employing PAUP, options (Mu with PRAP2 combination with in ana- performed ratchet SeqState were parsimony lyses trees, in parsimonious more implemented of & (Simmons as strategy (Mu (SIC) by 2000) informative coding as Ochoterena, indel coded either bisection- simple or were data a tree missing Gaps and as swapping. treated replicates branch 1000 reconnection with under sequence addition random searches using PAUP implemented Heuristic were using parsimony 2002). performed (MP) (Swofford, parsimony 4.0b10 were maximum criterion the optimality to according tions xn and exon, n fthe of end tal. et aiu ieiod(L nlsswsalso was analysis (ML) likelihood maximum A -) hs eeupdated. were these L-F), T T 4 4 le,20a.T erhtete pc o islands for space tree the search To 2004a). ¨ller, -trn -trn UGU UGU 08;pie C primer 2008b; , ńdez-Maqueda atwssltit w avs hc were which halves, two into split was part L (Herna L and eea ela the as well as gene ¨ller trn tal. et trn trn L L tal. et UAA ńdez-Maqueda rps T UAA 91,and 1991), , UGU 4 - -trn trn 06.Pyoeei reconstruc- Phylogenetic 2006). , 9 M - nrn hl h eodpart second the while intron, n fthe of end trn tal. et F trn eini opie none in comprised is region F trn yFrey by GAA L - (Herna L-F L UAA UAA 08) C products PCR 2008b). , trn atp pcr C protocols PCR spacer. H negncsaes the spacers, intergenic L tal. et tal. et B UAA intron, rps N la-pkit Clean-up DNA -rbc ee the gene, 4 08;primers 2008b; , 99 rmrF primer 1999; , ńdez-Maqueda Seh 2004; (Stech, L tal. et 5 9 le,2004b) ¨ller, xnad5 and exon trn 98.In 1998). , L UAA rps 4- 3 H 9 9 Published by Maney Publishing (c) British Bryological Society h Laayi,asnl pia rewsfound was tree optimal single In trees. L a shorter (ln analysis, find not ML did but the trees lengths recovered same searches the PRAP of 0.452]. 0.548, CI RC 4924, 0.620, lengths RI SIC: 0.523, with CI 0.326; RC 3325, trees, 0.623, RI lengths parsimonious indels: most [without two respectively or eight retained SIC total. parsimony-informative in 1186 characters parsimony-informative in 467 resulting another characters, were yielded positions) SIC variable characters by the indel of of Inclusion 3079 61.1% and parsimony-informative. or variable, comprised were (23.4 (38.2%) 719 alignment 1176 these, combined Of positions. The Brid. trn for (nt) 261–331 nucleotides were Dicranidae sequenced within the of markers ranges non-coding length dataset, present the In Results with runs Four (10 program. chains the four of settings default the The matrix. priori indel the the a employing for (‘F81’) partitions, model site unlinked treated restriction data and SIC indel separate by and as sequence coded with indels included, the were analysis, Bayesian second Rates 5 Rates 2.9656), Nst Basefreq 0.0845), cri- settings AIC the GTR the Consequently, and indicated test the terion Crandall, ratio Both 2005). likelihood & (Nuin, hierarchical (Posada MrMTgui 3.7 employing 1998) Modeltest in performed nueta nyteso h ttoaypaewere phase stationary the of trees included. only to that deleted were ensure (15%) trees 150 first trees). the sampled Consequently, 100–120 (first ‘burnin’, generations 000 000 120 100 the and first at the infer between runs ranged to all approximately which of and v1.5 (plateaus likelihoods) runs check comparable Tracer the to of used in convergence were for 2007) generated Drummond, & plots (Rambaut chains the Trace the after combining sampled trees converged. by the calculated using and were trees runs four clades consensus of rule PP majority 1000 and percent a every Fifty to written sampled file. were tree were trees respective heated Chains the single and 0.2. generations the to of set temperature chain the with neously, pcr olnt aito a bevdi the in observed nt the additional in one was for except variation regions coding sequenced length No spacer. for .74wr sdfrM nlssadNst and analysis ML for used were 0.2714 Paaye ihu n ihidl nlddby included indels with and without analyses MP - pcr 4–2 tfor nt 243–325 spacer, T-L trn 52 5 trn - pcr n 4–6 tfor nt 445–567 and spacer, L-F ngmafrteBysa nlss na In analysis. Bayesian the for invgamma rbblte upidwr hs pcfidin specified those were supplied probabilities 03.810,wihi hw nFgr 1, Figure in shown is which 20030.388180), L3 5 9 ,Rmat 6, xnin exon 5 6 am,Shape gamma, eeain ah eernsimulta- run were each) generations rps 5 aocpu nigritum Catoscopium 4- z 071 .66012 1.5770 0.1421 2.9656 (0.7917 trn I z pcr 5–2 tfor nt 252–328 spacer, T sbs-tmodel. best-fit as G trn 5 .59adPinvar and 1.3539 nrn 97 nt 59–78 intron, L 5 043 0.0722 (0.4531 atp (Hedw.) 5 B- and 6 rbc L o hw) u etdisd h ld ssse to sister as clade support. without the inside (tree only SIC nested Fissidens with substitutions but analysis on shown), Bayesian the based not in taxa and were 1) other or (Figure the analyses, to Bayesian sister example, for the relationships and For in analyses, contradictory. family also MP clades Other the some support in 1.00). high unsupported PP to remained moderate 78–80%, with (BS resolved of was relationship raceae close A support. Dicranaceae monophy- significant were with Ditrichaceae letic except families all clade, received unsupported taxa remaining one the BS(89%)inadditiontoaPPof1.00.Withinthislatter of formed clade clades third the these and 2), clade, of (Figure two SIC with first analysis the the In 1.00). (PP incl Pottiales remaining the third longirostris Eustichia as as Grim- Leucobryaceae second comprising the well 1.00), first PP BS, the (91% 1), miales (Figure analyses three the indels in latter, polytomy without a the as remaining resolved Within were the the clades 1.00). main by of (PP only clade analyses supported a Bayesian statistically to was sister which was species, and analysis, SIC 1.00) MP the PP in BS (84% next off branched Mitt. unsupported 2). an and phylogenetic 1 as the (Figures or clade in grades Dicranidae as the either of reconstructions, rest the to sister prorepens Drummondia Ochyra, nigritum Catoscopium support. significant Campylopodiella auso h epcieaaye ihSC The within showing SIC. only Leucobryaceae, with differs tree PP analyses the and parsimonious respective BS most showing the second 2, including of Figure trees in values parsimonious depicted most is two indels indicated. the indels of respective without the One analysis from Bayesian PP and and MP support bootstrap with fDcaia Seh 99,;L Farge La Goffinet 1999a,b; 2002; reconstructions (Stech, phylogenetic Dicranidae molecular of first the From first the (1999–2009) — decade Dicranidae of phylogeny Molecular Discussion ee rmodaee peeaee Erpodiaceae, Ephemeraceae, long Drummondiaceae, had as ceae, position Catoscopia- considered Amphidiaceae, (Archidiaceae, debated systematic been formerly whose monophyletic, or peristomes reduced diplolepideous, as either double with families resolved peculiar of number were a including Dicranidae First, Hedderson nalpyoeei eosrcin,tespecies the reconstructions, phylogenetic all In ircu flexicaule Ditrichum rhdu alternifolium Archidium nteM I nlss(iue2,albeit 2), (Figure analysis SIC MP the in tal. et s.str. ora fBryology of Journal tal. et 04,fu anrslsemerged. results main four 2004), , nsprt rnhs u without but branches, separate on yootaee n Calympe- and Hypodontiaceae, , 01 Tsubota 2001; , , tc tal. et Stech Bi. rd P .0,adthe and 1.00), (PP Brid. (Brid.) yeooacrispulum Hymenoloma roihu norvegicum Bryoxiphium Hd. .rto eeplaced were E.Britton (Hedw.) ddtx fDcaae and Dicranales of taxa uded (Schwa Amphidium hlgn fhpoeieu mosses haplolepideous of Phylogeny Atractylocarpus 2012 Hd. it and Mitt. (Hedw.) g. Hampe,¨gr.) and tal. et VOL 03 2004; 2003, , tal. et a placed was .34 (Hedw.) 2000, , (Brid.) and NO .3 175 Published by Maney Publishing (c) British Bryological Society tc tal. et Stech 176 ora fBryology of Journal iue1Snl pia aiu ieiodte f5 ersnaie fhpoeieu oss(irnde ae on based (Dicranidae) mosses haplolepideous of representatives 54 of tree likelihood ( maximum sequences optimal DNA Single chloroplast 1 Figure aiu asmn nlssadsgiiatpseirpoaiiis(P rmarsetv aeinaayi:BS analysis: respective Bayesian a respective from a values from (BS) (PP) support probabilities bootstrap posterior indicate significant . lines and Thick analysis representatives. parsimony outgroup maximum as used were (Encalyptidae) 5(lc) BS (black), 95 hlgn fhpoeieu mosses haplolepideous of Phylogeny . 70%/PP 2012 . 5(akge) PP grey), (dark 95 VOL rps .34 4 -trn region, F NO .3 . atp 5(ih grey). (light 95 B -rbc spacer). L imaaustriaca Timmia Tmide and (Timmiidae) nayt streptocarpa Encalypta . 90%/PP Published by Maney Publishing (c) British Bryological Society ancagsi hi eei opsto ( composition generic their and in ci changes traditional tain al- speciose their monophyletic in Fissidentaceae, more were most Pottiaceae, Calymperaceae, the and the Grimmiaceae, of as such some families, Second, Wardiaceae). and Splachnobryaceae, Rhachitheciaceae, Mitteniaceae, ea reat h ooyi itnaeewere Mitteniaceae monotypic The artefact. an be of Goffinet phylogenetic position in Bryaceae the diplolepideous in The indicated not tree. and in given appendix only were the names taxon the as possible article not that from was however, systematic taxa, the individual of on position Inference recon- genera. phylogenetic moss large-scale of a struction on based mosses in included Shaw below), in only Pleuro- were to monogeneric Serpotortellaceae (see and the difficult phascaceae Pottiaceae of remained Representatives family Pottiales assess. of large circumscription the relationships within resolve to efforts Despite monophyletic. Maqueda (Tsubota well-resolved relationships molecularly with group monophyletic a represent to shown been unequivocally have Seli- geriaceae) and Ptychomitriaceae, Grimmiaceae, steliaceae, (Campylo- Grimmiales only Pottiales, and Dicranales, Goffinet in not but & Frey (2009), in Stech incorporated were which 2008), Frey, & Dicranellaceae (Stech families and described newly Dicranaceae Aongstroemiaceae, or Amphidiaceae, resurrected of the into segregates placed Further ( 2009). genera ‘dicranoid’ both and comprising heterogeneous ‘leucobryoid’ Leucobryaceae more the a of in circumscription resulted which Dicranaceae, Paraleucobryoideae and and Pottiaceae, Campylopoideae subfamilies the former the by in expanded were Leucobryaceae included were Ephemeraceae Dicranaceae the in and included family new Dicranaceae a the from as separated (Rhabdoweisiaceae) Pottiaceae Oncophoraceae the from segregated Stech, recent & Goffinet main (Frey two Bryophyta 2009; of the classification in of below). incorporated synopses and enumeration rearrangements made systematic (see were of phylogeny number the a Subsequently, in first off diverse (Hedderson taxa ‘proto- morphologically haplolepideous’ called families, different a from species Fourth, of assembly phylogeny. the the and in recognizable supported not was weakly Dicranidae especially of rather classification ordinal was polyphyletic, of backbone phylogeny the as the Third, Ditrichaceae. resolved and Dicranaceae were contrast, Herna ftetrelretodr,nml Grimmiales, namely orders, largest three the Of ńdez-Maqueda Jaffueliobryum tal. et tal. et s.l. 20) h sesdmlclrdiversity molecular assessed who (2005), 08) irnlswr lal not clearly were Dicranales 2008b). , utemr,Dceoaeewr re- were Dicnemonaceae Furthermore, tal. et tal. et 09.Hpdnica were Hypodontiaceae 2009). , eoe rmGrimmiaceae; from removed tal. et Pleurophascum 08) te aiis in families, Other 2008b). , cmcito rwt cer- with or rcumscription s.str. 03 Herna 2003; , tal. et tal. et icioaeeand Cinclidotaceae , tal. et cf. 04,branched 2004), , (2009). ry&Stech, & Frey 20)might (2001) ihnthe within p.p. Indusiella s.l. ńdez- fthe of were rd ruspotdcaei h phylogenies, the in clade (Hedderson Leucobryaceae Archidiaceae, unsupported Grimmiales, or comprises as grade appears which group, a first The groups. large two into Tsubota Farge (La tions to close Milde of position the data. molecular by s.str. supported view of al. et ihricue nPtils(Goffinet Pottiales in included either ry&Seh 2009; Stech, 1985; (Shaw, & Mitteniales order Frey separate a as treated ags oto ftehpoeieu oss namely mosses, haplolepideous the of the portion comprises group largest second The study). (this chiaceae (Cox glaucescens (Goffinet Saelania Micromitriaceae described otaee(Cox Pottiaceae tasmanicum Tridontium to sister Timmiella as (Ditrichaceae) Reimers of position the study), nlddin included study. further h allpdosmse opieCox Wahrmund comprise mosses haplolepideous within the relationships of reconstruc- inferences phylogenetic further allowing molecular tions recent most The state current recent and — developments Dicranidae of phylogeny Molecular icsini tc ry 2008). Frey, & Stech in discussion 00 Wahrmund 2010; cip)Lmr Ptica,seblw.Teclose crispulum Hymenoloma The between below). see relationship (Pottiaceae, Limpr. Schimp.) and (Ditrichaceae), crispulum enepne yincluding has by Dicranidae expanded of been circumscription The study. present cuei aquatica Scouleria (Drummondi sum- in Scouleriales comprise, includ- to Distichiaceae, Catoscopiaceae, seem study Bryoxiphiaceae, they every mary, taxa, not respective Although all Cox ed 2). and in a 1 (Figures such resolved as also real, Wahrmund be is to of topology seems the number in phylogeny, a first off Dicranidae of branching of family existence taxa findings & diverse the surprising morphologically as studies, Frey most earlier the by separated of several One Hedwigiales already (2008). the Stech was within Bryowijkiaceae which 2010), h te allpdostx emt edivided be to seem taxa haplolepideous other The 20)bsdo opooia hrces point a characters, morphological on based (2003) lal eog oRadwiica codn to according Rhabdoweisiaceae to belongs clearly tal. et tal. et (Cox (Rhabdoweisiaceae), 00 Goffinet 2010; , tal. et tal. et Dicranoweisia tal. et 2004). , Rhabdoweisia yeooacrispulum Hymenoloma irnwii cirrata Dicranoweisia ora fBryology of Journal tal. et 04 tc ry 08 Cox 2008; Frey, & Stech 2004; , tal. et 21) Goffinet (2010), tal. et tal. et ok)a elas well as Hook.) 21) n h rsn study present the and (2010), 00 Goffinet 2010; , tc tal. et Stech Hd. rt.o Ditrichaceae of Broth. (Hedw.) imel anomala Timmiella 02 Hedderson 2002; , hyolsel chilensis Chrysoblastella 00,adtepsto of position the and 2010), , (Hedderson 00 swl sterecently the as well as 2010) , rmodaprorepens Drummondia cf. okf Solraee in (Scouleriaceae) Hook.f. u eaae yOchyra by separated but , ca,Scouleriaceae aceae, npyoeei reconstruc- phylogenetic in loOBin 07and 2007 O’Brien, also tal. et Bryowijkia hlgn fhpoeieu mosses haplolepideous of Phylogeny ircu flexicaule Ditrichum tal. et 2012 01,adEusti- and 2011), , tal. et Hd. id.ex Lindb. (Hedw.) tal. et a traditionally was Distichium tal. et 21) n the and (2011), tal. et tal. et 04 present 2004; , Dicranoweisia Hymenoloma VOL tal. et tal. et (Cox 01 need 2011) , Buh& (Bruch 09 or 2009) , .34 2011), , (Mont.) ,2004; (2010), (2010), tal. et tal. et p.p. NO and and .3 , : , 177 Published by Maney Publishing (c) British Bryological Society tc tal. et Stech 178 ora fBryology of Journal aeinaayi ihidl nlddaegvna h branches. the at given are included indels with analysis Bayesian ugoprpeettvs otta upr values haplolepideous support of representatives Bootstrap 54 representatives. of included. phylogram) outgroup (SIC) as coding ( (shown indel sequences reconstructions simple DNA parsimonious by chloroplast most on two based of (Dicranidae) out mosses One 2 Figure hlgn fhpoeieu mosses haplolepideous of Phylogeny 2012 VOL .34 imaaustriaca Timmia NO .3 . Tmide and (Timmiidae) 0 n infcn otro probabilities posterior significant and 70% rps 4 -trn region, F nayt streptocarpa Encalypta atp B -rbc pcr,wt ne hrcescoded characters indel with spacer), L Eclpia)wr sdas used were (Encalyptidae) . 5fo respective a from 95 Published by Maney Publishing (c) British Bryological Society flexicaule Trichodon E.Britton, between (Hedw.) pallidum indicated Ditrichum is ship Goffinet and with clade ported into reconstruc- cluster phylogenetic tions. molecular to the seem in far groups two so analysed Ditrichaceae well of as study present 2). the Figure versus 1 in (Figure ambiguous remains Goffinet Goffinet Hedderson 2004; excluding significant and, 2) receives Amphidium (Figure indels group with analysis present The this Wahrmund Pottiales. in speciose of support statistical most clade the as respective well as kiaceae Bryowij- plus Schistostegaceae) and Rhachitheciaceae, Oncophoraceae, Hypodontiaceae, Fissidentaceae, ceae, Ditrichaceae Dicranellaceae, Dic- ranaceae, Calymperaceae, Bruchiaceae, (Aongstroemiaceae, Dicranales of families most ecmltl ovdwt rebsdapoce (e.g. approaches tree-based Hallstro with some not solved might in completely relationships be Although level higher study. lineages further organism need pre- as and remain Ditrichaceae, liminary such especially families and phoraceae, of relationships Onco- Dicranellaceae, Bruchiaceae, Aongstroemiaceae, and circum- discussed Besides, scriptions phylogenies. be extended such to on needs Bell based by lineages, e.g. level, pleurocarpous done, main family been the has above as lineages to or names major useful node-based characterize formal informal remain by will replaced of be Dicranidae should distinction the within ordinal the orders an preliminary versus Whether the groups classification. major concerns three this of level, distinction levels. highest taxonomic the different At at relationships Dicranidae ambiguous the still within clarify sampling to marker and necessary taxon are increased on ana- based molecular lyses phyloge- of further However, reflects classification relationships. better a netic that towards mosses step haplolepideous important an repre- them sent from inferred material. rearrangements systematic further 1 by (Figures confirmed study be to present needs the 2) of and trees the in position to close might group Ditrichaceae latter This represent lacking. is support statistical xetfor Except h vial hlgntcrcntutosadthe and reconstructions phylogenetic available The Cladophascum Eccremidium m&Jne 00,tercn eeomnsof developments recent the 2010), Janke, & ¨m and , tal. et tal. et Ceratodon L Farge (La tal. et Apiica) nCox in (Amphidiaceae), 01.I otat ls relation- close a contrast, In 2011). , alnaglaucescens Saelania hyolsel chilensis Chrysoblastella 21) h oiinof position The (2011). 01 rsn td) although study), present 2011; , Aongstroemia tal. et Pleuridium Hd. Hampe/ (Hedw.) Buhaee (Cox (Bruchiaceae) tal. et and swl MDne,20) its 2005); (McDaniel, well as .t.Cheilothela s.str. 02 Tsubota 2002; , 04 Cox 2004; , Garckea sensu Astomiopsis , Pseudephemerum tal. et (Aongstroemiaceae) ry&Seh(2009) Stech & Frey tal. et omawell-sup- a form tal. et representatives , .heteromallum D. 20)frthe for (2007) p.p. 21) nthe in (2010), tal. et , , tal. et tal. et Amphidium 21)and (2010) Erpodia- , hudbe should Ceratodon Ditrichum 2010; , 2010; , 2003, , ,and , irndewt o-oigmresmgtb the be might markers non-coding of with phylogeny backbone Dicranidae the [only resolving of 2004 problem One (Stech, mosses trn family haplolepideous at in the successfully level employed of already spacers were which by replaced was nomto,a h eyshort present very phylogenetic the the analyses. more (2008), as MP Frey provides information, the & combination in Stech marker least to at Compared lar- unsupported, remain well relationships gely very of their families are whereas different study, the Dicranidae, support present and resolve the to suited in sampling. (2008) especially Frey taxon & and Stech larger in a tested as with markers, Non-coding employ thus to not are useful and did not Dicranidae rates, within evolutionary relationships moderate resolve to slow show adpatrltosisb Qiu by under infer relationships to level used plant markers land coding taxonomic the example, respective For study. phylogenetic the and at variability identified structure be sufficient must markers provide comparing new that time and same the at markers continued. But be should multiple genomes different from using markers of the (2010) general, in trend Quandt bryophytes for & in discussed recently Stech uncertainties have As remaining phylogeny? the Dicranidae resolve to followed study present mosses, Dicranidae. the the in of for extended and plastid reconstruction continued is non-coding phylogeny which combined for evaluat- of who markers suitability (2008), Frey the & ed Stech by followed was ye eebsdo he ieyue lsi DNA plastid used widely three regions, on based ana- were phylogenetic lyses higher-level initial all Dicranidae, molecular few only ( on markers focus in the analyses been phylogenetic has many bryophytes of problem general on A based sampling marker made Molecular sets. be data extended still of reconstructions can phylogenetic progress considerable hlgntcrsac nteDcaia niaethat indicate Dicranidae the in research phylogenetic tal. et (Qiu genomes plant Quandt even different or two three from all markers the combined used from and (Wahrmund 2010), especially genome plants) markers, land mitochondrial new all evaluated (or mosses have of phyloge- reconstructions improve genome. netic only to plastid approaches thus the recent Dicranidae of Fortunately, patterns of evolutionary analyses reflect initial all 2010), ihsnl-akraaye eg lwnk Page, & Gontcharov Slowinski 1999; (e.g. analyses single-marker with hc taeyo akrslcinsol be should selection marker of strategy Which - pcr;Herna spacer]; T-L 00 Goffinet 2010; , trn tal. et cf. L - tc unt 00.Cnenn the Concerning 2010). Quandt, & Stech F, 07 Cox 2007; , ora fBryology of Journal rps ,and 4, tal. et tc tal. et Stech ńdez-Maqueda tal. et 04 el&Hyvo & Bell 2004; , rbc 01.Aohrstrategy Another 2011). , tal. et .Aiefo problems from Aside L. hlgn fhpoeieu mosses haplolepideous of Phylogeny tal. et 2012 00 Wahrmund 2010; , psb rps A 4 tal. et tal. et 20) which (2006), -trn -trn VOL tal. et .34 region, L spacer H 2008b). , ,2009, 2006; , ¨nen, NO .3 179 Published by Maney Publishing (c) British Bryological Society tc tal. et Stech 180 ora fBryology of Journal 05 swl as well as 2005) led tlzda oe aooi ee in level taxonomic lower at are Dicranidae utilized already markers. nuclear markers, and organelle the nuclear between evolutionary patterns low-copy different potentially account or into taking newly single- and should markers developed markers mitochondrial with plastid combined suitable be most The tested. be rdcd od o ute lsi akr uhas such ( markers introns plastid 2 further be group so, can char- do trees To indel phylogenetic produced. considering well-resolved also until (thereby acters) them in sites of synapomorphic of each amount markers small suboptimal the collect several to combine to has perhaps some study. for present support the higher in provide clades, even substitu- and the data, with tion congruent these generally regions, are DNA SIC. high characters non-coding the of by given variability indel levels length use coded taxonomic to higher critical indels at considered characters be the might it the in Although in higher than even be substitutions to be seems present homoplasy the of can amount of ( values reconstructions as RC parsimonious low most homoplasy, the from e.g. of inferred, amount considerable sn prahslk hs ulndi Tekle in outlined those below), like (see data approaches genomic using available of on based comparison set characteristics a a appropriate identify to with effort markers concerted of A gene analysis. complex- the of the ity and increase dramatically contain micro-inversions, that may duplications nrITS, deletions, as such insertions, evolving, freely too multiple that are markers experienced However, phylogeny. already the across have changes sites change few the can because that misleading functional be rigid may under constraints, are they loci because are or they of genes because Analyses small either sites, challenging. variable few the more contain is however, that loci analysis, of of the level choice of this backbone For the for Dicranidae. resolving particularly equally like be phylogenetics, problems to thorny deep likely design multi-copy for are a to genes important of Nuclear paralog straightforward family. single gene a relatively duplicate amplify of to is primers portions it coding the close genes, than for faster diverge introns much explanations Because distinguishing species. among as relationships for polyploidy) critical hybridization (including and sorting lineage are closely incomplete between loci distinguish multiple, second, and independent to populations; or variable species related are sufficiently introns nuclear First, often reasons. two for scale analyses this genetic at evolutionary for essential are introns n ateovn ee uhas such genes fast-evolving and xmlso ige rlwcp ula markers nuclear low-copy or single- of Examples sSeh&Qad 21)frhrdsusd one discussed, further (2010) Quandt & Stech As hlgn fhpoeieu mosses haplolepideous of Phylogeny adk trn 2012 gpd G, and trn Wl,20,20) Nuclear 2005). 2002, (Wall, VOL V, phy .34 rpl MDne Shaw, & (McDaniel 2 mat 6 and 16, NO Kor .3 cf. trn ndh eut) The results). introns) K ,should B, tal. et h aeo iesfiaina h aeo the of be base may the loci at additional required. that diversification indicates the of but Dicranidae candidate, rate that strong a the suggest is family gene data) phytochrome unpublished analyses Preliminary (McDaniel, forward. way a provides (2010), emn ftePL(n-ihho h peristome) (Shaw the develop- of mentally peristome haplolepideous (one-eighth two-cell the PPL a characterizes for the cells of 2:3 of segment arrangement PPL:IPL leading a divisions to cell of sequence (Shaw unique a types Thereafter, peristome and major peristome other the haplolepideous the primary virtually between are (OPL), identical (IPL), layer outer peristomial inner the and differentiated namely (PPL), is layers, amphithecium three the into where point the 1957). Anderson, characters & (Bryan sporophyte in genera differences both and between known long numbers the chromosome (Goffinet to addition unrelated in 2011), molecularly are which the reductions is (in peristome taxa example, including diverging considerable, first For already the char- Dicranidae. of diversity of in morphological interpretation evolution the for acter challenges that morphologies pose different will very major here with distinguished taxa three comprise mosses All haplolepideous coding. of further groups character supported homology need and better for also assessment and probably analyses expanded will morphological–anatomical on and as based phylogenies need well turn, be in as cladistic to These, reconstructions. characters on state morphological based ancestral of be should analyses into evolution insights precise character More morphological haplole- below. the key peristome for preliminary pideous of discussed exemplarily allow evolution as characters, the data of molecular inferences available The evolution character for Implications atr f8:4: (Este of final a and pattern 1979), (Edwards, Hypodontiaceae in 2(–3) PPL and by displayed is reduction gametophyte lel of in the ture as such haplolepids, p.p. Scouleria : atr.Rsligfrua r,eg 4: e.g. a (4:) to are, Seligeriaceae, reduction formulas or Resulting teeth pattern. of the 2:2 row to second due a taxa of haplolepideous formation several in however, modified peristome), is mature the in disappeared have h al tgso eitm eeomn,u to up development, peristome of stages early The Timmiella ´banez Bryoxiphium IL4: :IPL Micromitrium Bryoxiphium 2 tal. et 3(r0: (or :3 seblw.Asrkn xml fparal- of example striking A below). (see n iia opooiswt other with morphologies similar and ) , 2 tal. et 2 06.Dul haplolepideous Double 2006). , 2–)i ayprca,(8:4:) Calymperaceae, in :2(–3) Catoscopium n h otaeu morphology pottiaceous the and in ftefre Ephemeraceae, former the of 2 3a h P/ue P walls PPL OPL/outer the as :3 99.TefruaOPL: formula The 1989). , lpoiru humillimum Glyphomitrium Fissidens , Drummondia lk efarchitec- leaf -like tal. et Ephemerum 2 2–)in :2(–3) 1989). , tal. et ,and 2 , : Published by Maney Publishing (c) British Bryological Society eitm of peristome on the Calymperaceaediscussion for well-supported and synapomorphic (except syrrhopodontoid be and to The monophyletic seem 2009). types Stech, pottioid ( & type pottioid Frey dicranoid, and the syrrhopodontoid, namely types, seligerioid, main four into grouped several teeth. Dicranidae. whole peristome occurred the across the obviously independently of times reductions surfaces of outer Peristome ornamentation and and incision, inner of degree to shape, (Shaw, respect the with layer considerable furthermore is endothecial peristome haplolepideous the outermost of variation Morphological the cell 1985). adjacent and of IPL the walls of to walls restricted periclinal ( inner usually rarely the or is teeth, the which of side, base the OPL forma- the preperistome on from tion result mostly peristomes f8o 6etr et,wt h formula the with teeth, entire 16 or 8 of swl si h oeual distant molecularly the in Werner by dataset as (molecular well Pottiaceae of genera as different have in to times seemed several membrane, evolved basal a from arising teeth by Ellis, 1990; Eddy, (e.g. 2007). Octoblepharaceae family Octoblepharum (Tsubota (Tsubota support Hedderson (well-supported) 2003; low remaining with the to Calymperaceae In sister as 1979). resolved (Edwards, phylogenies, inferences makes difficult molecular which relationships tooth, single about a to referring allpdoslnae (Cox Onco- lineages haplolepideous and Rhachitheciaceae, ( phoraceae Grimmiales, (other) riales, in investigated. peristomes be to twisted remains the Pottiaceae to homologous of development deep indepen- the the on Whether shared not evolution. parallel peristome), dent twisted (a of a with homology peristome ancestor developmental twisted ‘deep’ the interpretation, Timmiella this lineage. silenced In phylogenetic be another in can later evolved, re-activated gene once and and, a adaptive that highly i.e. organs is major of homoiology, existence the of determining cluster example an of represents morpho- other traits to argued similar logical (2006) peristome, twisted Zander the However, that 2006). Zander, by eetees allpdosprsoe a be can peristomes haplolepideous Nevertheless, h xrsino h otodprsoedisplayed peristome pottioid the of expression The aawt eieii eitmsocri Scoule- in occur peristomes seligerioid with Taxa Timmiella Timmiella Octoblepharum tal. et n ohr otaeersle rma from resulted Pottiaceae (other) and Glyphomitrium ih3 lmnos pcls,twisted spiculose, filamentous, 32 with , Octoblepharum eitm si atdevelopmentally fact in is peristome Timmiella 04.Teerslsidct that indicate results These 2004). , ih ebte lcdi t own its in placed better be might tal. et Timmiella 04 rsprtdfo them from separated or 2004) , n otaee(u see (but Pottiaceae and ) Octoblepharum ,wihbln odifferent to belong which ), eo) epciey The respectively. below), speimrhcand plesiomorphic is , srdcdadconsists and reduced is tal. et Mittenia tal. et 04 re-analysed 2004; , 00 Goffinet 2010; , yinvolving by ) a either was Timmiella 2( tal. et 2 3) cf. :2 , tal. et Maqueda the of genera, analyses largest phylogenetic with well-studied, & already quite are Stech Grimmiaceae within 2004; Relationships 2010). with Stech Dohrmann, 2006; Stech, & & deal Frahm 2005; Wagner, Stech which 2004; of (Stech, six phylogenetic molecular detailed publications, more 18 only by ered te ag allpdosgnr,sc as such genera, relationships. haplolepideous large and Other circumscriptions taxon assessing s.l. h n f21) seilywt epc olarge to (100 respect genera largest with 11 haplolepideous The Especially understudied. seem 2010). Dicranidae genera, of end to up publications the by extended 2010; in Quandt, compiled & (literature Stech taxa haplolepideous with (22%) 65 deal only mosses, 292 on of studies total systematic a molecular of moss Out underrepresented. however, be total to record, seem publication they the the species. of In single diversity. 30% species or about genera comprise within relation- Dicranidae and biogeographic so between and published ships systematic studies tackled most systematic far mosses, higher-level in on relationships focussed era’ ‘molecular the of beginning the in analyses phylogenetic Although phylogenetics Genus-level fdcaodadslgrodprsoetpsare (e.g. Shaw types studies 1979; earlier peristome Edwards, complement diversity seligerioid to the necessary and clearly covering dicranoid taxa selected of of analyses developmental and histochemical, compara- morphological, Further tive Grimmiaceae. distribution, seemed the phenolics) characterize family, of (pectin to quantity same maximum properties the with can stages of certain properties species histochemical although between to greatly by relation vary in peristomes ment seligerioid in mainly especially Este of type, putatively comparison dicranoid peristomes ( peristome the of Ditrichaceae the presence from of the surfaces reduced outer and and teeth, inner ornamenta- and of incision of tion degree the peristomes to dicranoid respect with of especially by variation supported large represent is the hypothesis latter could The morphologies. peristome evolved or non-related different have of independently, assemblies thus artificial could times types several dicranoid and ioid Oncophoraceae and Fissidentaceae, ytmtcrlvneo hrcesi hs peristome these in types. characters of relevance systematic tmsaefudi eea ifrn aiissc as such families different Ditrichaceae several in found are stomes al. et ´banez (Larraı 09 Milyutina 2009; , 01.Smlry aawt irni peri- dicranoid with taxa Similarly, 2011). , tal. et ń tal. et p.p. tal. et cf. ora fBryology of Journal 07 2008a,b), 2007, , rmi s.l. Grimmia 20)soe htprsoemove- peristome that showed (2002) ecbyca,Dicranaceae Leucobryaceae, , ry&Seh 09.Bsds the Besides, 2009). Stech, & Frey 01,poiigabssfor basis a providing 2011), , tc tal. et Stech tal. et tal. et Srif 06 Herna 2006; (Streiff, z 00,and 2010), , 99 n oifrthe infer to and 1989) , pce ah r cov- are each) species hlgn fhpoeieu mosses haplolepideous of Phylogeny Schistidium 2012 p.p. Racomitrium VOL h seliger- The Campylopus tal. et .34 (Ignatova Fissidens ńdez- 2007, , s.str. NO .3 , 181 Published by Maney Publishing (c) British Bryological Society tc tal. et Stech 182 ora fBryology of Journal rdtoa eei eiiain,freapebetween contradict example cases Weissia for many delimitations, is in generic the data traditional Trichostomoideae On molecular levels. hand, taxonomic subfamily one all at The complex particularly 2003). 2002, Syntrichia subulata Tortula (e.g. large genera several difficult Didymodon includes taxonomically it because & and (Frey and genera 83 2009), in Stech, species moss 1425 largest about the with is family, it because difficult particularly are therein). references ( Kuc & Werner e.g. sequences in, published few included The unavailable. almost are as such genera Trichostomum (Stech data molecular and circumscriptions level species morphological molecular between incongruence the as such genera at unknown (Werner almost remain eune nsmlsta oeatosseparated authors e.g. some genera, that different identical into samples almost show in nrITS evolving sequences like rapidly markers Even variation. molecular the molecular with of compared degree as morphologically fast genus the extremely hand, other the On aete eeabsdo diinlmressol eper- be should contrast, markers In additional formed. on based genera further of topologies, analyses molecular phylogenetic molecular such both treat to of how of species tions by formed Crossidium being clades supported s.l. Pterygoneurum of the members in included traditionally species genus the of part a that of circumscription the Tortula is (Werner complex data Especially nrITS 2005a). by supported char- not is morphological like acters on based genera taxonomy Other subgeneric needed. clearly Didymodon is molecular–morpholo- analysis combined gical a but polyphyletic, ee eg Werner Po (e.g. level of relationships resolve to al. et and Kindb. (Limpr.) eaae from separated tal. et c 4 p. or spp.) 440 . naeln eainhp ihnPottiaceae within relationships Unravelling hlgn fhpoeieu mosses haplolepideous of Phylogeny , ld (Werner clade 06 adr 06,amr opeeanalysis complete more a 2006), Zander, 2006; , 05) nsmay ept oeua efforts molecular despite summary, In 2005b). , Oxystegus ˇ r 21) emt niaethat indicate to seem (2011), era Pottia , hl h vial aaspotteview the support data available the While . tal. et Trichostomum namrhlgclbss(Werner basis morphological a on , and emt emnpyei,bttheir but monophyletic, be to seem Hyophila 09.Hwvr loqiewell-studied quite also However, 2009). , ,and r nedmrhlgclyreduced morphologically indeed are Tortula and , Campylopus , Barbula Tortula Dicranella Chionoloma tal. et Tortula ew hwsm iiaiywith similarity some show Hedw. 2012 tal. et 02 04 05,;Grundmann 2005a,b; 2004, 2002, , Syntrichia Stegonia Weissia .controversa W. , ,also , sd rmgnrlconsidera- general from Aside . and lhuhsm pce like species some although , Syntrichia Pottiopsis tal. et 02,wt eea well- several with 2002), , VOL / Leptotrichella Weissia eanacalnedeto due challenge a remain ,and Hyophila .34 taeea (supra-)generic at ttiaceae .Frsm fthese of some For ). Weissia Crossidium 20)adKo and (2004) r ato a of part are smlclryclearly molecularly is , Pseudosymblepharis , Tortula NO Tortella Tortella [ Astomum .3 oeua data molecular , tal. et ew (Werner Hedw. em oevolve to seems ( , c n related and , 00and 2010 , 2 spp.) 220 . Barbula , Phascum Pleuroch- ¨ckinger Barbula Tortula ] Tortula tal. et levieri tal. et s.str. is , , , . , , eti h rgn rmacosbtena between cross a of from population eus tropical develop- progeny a abnormal and the temperate and in architec- inviability genetic ment spore the dissect of to ture map genetic a used rmica rvee nNthv Cronberg, McDaniel recently, & More Natcheva 2004). in (reviewed and Grimmiaceae sporophyte Dicranaceae, the Pottiaceae, in Ditrichaceae, patterns hybrid families germination with spore documenting mosses, and morphology haplolepideous studies the several in history long in species purpureus moss a in among-population variation and genetic quantitative within in- of most Beer description the & depth conducted Shaw (2005) and McDaniel and soils, (1999) containing metal heavy on that demonstrated via populat natural targeting with work gene Trouiller efficient (Brucker recombination undergo homologous to known and yeast than other eukaryote, only Besides, 2006). Quatrano, & Cove (Bowker variation ratio sex al. on et research for model a (Oliver al. sequenced et was after genome chloroplast species, complete moss second patens the P. is D.Mohr & Weber Syntrichia in haplolepideous on structure namely focussed taxa, mostly genomic far so and has mosses mechanisms genetic on Beike in (reviewed (Funariidae) moss’ patens model ‘genetic Physcomitrella prominent most the from Apart genomics and Genetics h oa iest ihnPtica ssilinsuffi- still is Pottiaceae known. that within ciently indicating diversity 2011), total Hedderson, the & Zander 2011; Ko 2010; eeomna epne olgtadgaiy(Cove of gravity study and the al. for et light model to a responses as used developmental condi- widely is laboratory and under tions, growth and mutagenesis to Oliver (e.g. Stark tolerance desiccation and 08,;Gleo&Cn,20,20;Erdag 2009; 2007, Cano, Ku & Jime Gallego 2008a,b; 2008a,b; 2007, & (Hedderson Zander, data on morphological based and/or described new molecular recently were remarkable genera several and Besides, species genera, missing. sampl- still larger marker is and the ing, taxon comprehensive of a on especially based and Pottiaceae, of rcnr 09 Cano 2009; ¨rschner, nraigterslto ftepyoeyof phylogeny the of resolution the Increasing . 00,and 2010), , 96 Sack 1996; , 00,sxa iopim(Stark dimorphism sexual 2000), , tal. et ¨ckinger tal. et swl.Tesuyo yrdzto a a has hybridization of study The well. as (Sugiura 2006). , species. eaoo purpureus Ceratodon 07 Mittmann 2007; , Syntrichia tal. et tal. et .purpureus C. eaoo purpureus Ceratodon ytiharuralis Syntrichia tal. et 00 kym Goffinet, & Akiyama 2010; , 01 Thornton 2001; , Hd. rc Schimp. & Bruch (Hedw.) tal. et 03,fo hc the which from 2003), , os ue hw(1994) Shaw & Jules ions, pce r elkonas well-known are species 00 Jime 2010; , nz&Cn,2007, Cano, & ńez tal. et a dp ogrowth to adapt can tal. et .purpureus C. Hd. rd and Brid. (Hedw.) tal. et .patens P. 00,research 2010), , tal. et 20,2008) (2007, tal. et al. et Hd. F. (Hedw.) samenable is tal. et 09.In 2009). , ńez .purpur- C. 2001), , 2005; , htis that , 2005; , 2005; , tal. et sthe is ˘& C. , Published by Maney Publishing (c) British Bryological Society o,CJ,Gfie,B,Wcet .. oe,SB hw A.J. Shaw, & S.B. Boles, N.J., Wickett, B., Goffinet, C.J., Cox, dy .1990. A. Eddy, 2006. R.S. Quatrano, & D.J. Cove, hag .. cal ..&Pn,CI 1998. C.I. Peng, & B.A. Schaal, T.Y., Chiang, Jime M.J., Cano, Este Erdag li,LT 2007. L.T. Ellis, 1979. S.R. Edwards, 1996. E. Hartmann, & R.S. Quatrano, D.J., Cove, rce,G,Mtmn,F,Hrmn,E apre,T 2005. T. Lamparter, & E. Hartmann, F., Mittmann, G., Brucker, 2000. B.D. Mishler, & D.N. McLetchie, L.R., Stark, M.A., Bowker, ra,VS nesn ..1957. L.E. Anderson, & V.S. Bryan, Vervliet- D., Hyvo Lang, & W., N.E. Bell, Frank, E.L., Decker, A.K., Beike, el .. unt . ’re,TJ etn ..2007. A.E. Newton, & T.J. O’Brien, D., Quandt, N.E., Bell, kym,H&Gfie,B 2011. B. Goffinet, & H Akiyama, References Nil. Changes: and Additions assistance. to Taxonomic technical for due (Leiden) Eurlings are M. C. grant thanks M. Spain of a Sincere by Technology supported CGL2009-09530-BOS. by and is Science JM possible of RHM. Ministry made to grant was SYNTHESYS sequencing Additional Acknowledgements genome critical and a group. family this provide across gene evolution as studying well for or framework as developmental studies, from population derived correlations to acter biologists test evolutionary sophisticated develop enable will Dicranidae bnz . sbt,H,Ymgci .&Dgci .2002. H. Deguchi, & T. Yamaguchi, H., Tsubota, B., ´banez, genera. 2010. cdmaSinica Academia between spacer and noncoding a sequencing and amplification yi eu rmSuhAmerica. South from genus typic microdonta Guerramontesia uky ihsm eak nteseito eteo the of centre speciation the on remarks genus. some with Turkey, Ku Octoblepharaceae. 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Wall, enr . o,RM,Cn,MJ ura .2003. J. Guerra, & M.J. Cano, R.M., Ros, O., Werner, Jime O., Werner, enr . o,RM,Cn,MJ ura .2004. J. Guerra, & M.J. Cano, R.M., Ros, O., Werner, 2002. J. Guerra, & M.J. Cano, R.M., Ros, O., Werner, enr . Patin O., Werner, herb. nn364()JQ690736/AF135101 (B) 36664 onen hnra N n w hools loci. mito- chloroplast non-coding Evolution two and of and types DNA three chondrial in information 2009. phylogenetic V. Knoop, taxon. otaee(roht)fo ot fia n nertmfor erratum an and Acaulonopsis Africa, South from (Bryophyta) Pottiaceae 602. data. sequence nrITS based Bryophyta) on (Pottiaceae, Trichostomoideae of phylogeny eunedata. sequence eetydvre iegsin lineages of reconstruction diverged phylogeny for recently dehydrogenase phosphate 10–26. 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F hlgn fhpoeieu mosses haplolepideous of Phylogeny Picobryum 2012 Mitthyridium and EU186590 EU186592 JQ690701 JQ690702 rps sequences. 4 ln Systematics Plant ln Systematics Plant otl bolanderi Tortula e eu of genus new a , VOL 4:147–64. 243: , atp .34 Tortula 0:581– 100: , Molecular Molecular B Fissidens -rbc (Musci: nthe On Nova 25: , NO rps and L .3 4 185 Published by Maney Publishing (c) British Bryological Society tc tal. et Stech 186 ora fBryology of Journal ecbymjuniperoideum Leucobryum denudatum Dicranodontium flexuosus Campylopus Isoviita & J.-P.Frahm flagellacea Campylopodiella imaaustriaca Timmia (outgroup) Timmiaceae acuta Blindia Seligeriaceae polyphyllum Ptychomitrium Ptychomitriaceae muralis Tortula ruralis Syntrichia riparius Cinclidotus Pottiaceae crenulata Rhabdoweisia bogotensis Oreoweisia wahlenbergii Oncophorus virens Oncophorus crispulum Hymenoloma polycarpum Cynodontium Oncophoraceae africanus Pilopogon gardneri Ochrobryum leana Brothera alticaulis Atractylocarpus Leucobryaceae pomiforme Hypodontium dregei Hypodontium Hypodontiaceae apocarpum Schistidium canescens Racomitrium Grimmiaceae fontanus Fissidens bryoides Fissidens Fissidentaceae longirostris Eustichia Eustichiaceae biseriatum Erpodium Aulacopilum Erpodiaceae prorepens Drummondia Drummondiaceae pce oce c.no. 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