is supported in the ntall (75% BS), ntno3rd (70% BS), and AA (78% BS) analyses. In the RY analysis, the relationship Tracheophytes (Vascular Plants)

Transkriptio

1 : iso kuva Yleiskuva Kasvisystematiikka: maakasvit Jari Oksanen Ekologian yksikkö Jari Oksanen (Ekologian yksikkö) 1 / 43 Plastidigeenit Ruhfel et al. BMC Evolutionary Biology 2014, 14:23 Page 7 of 26 Superrosidae Eudicotyledoneae Pentapetalae 69 Yleiskuva Ruhfel et al BMC Evolutionary Biology, 14:23 Superasteridae Malvidae Fabidae Lami. Campanuli. 99 Angiospermae Mesangiospermae Spermatophyta Embryophyta Tracheophyta 52 Streptophyta Monocotyledoneae Commelinidae Magnoliidae Acrogymnospermae Gnetophyta Lyco. Monilophyta Chlorophyta Brassicales Malvales Sapindales Crossosomatales Myrtales Geraniales Rosales Cucurbitales Fagales Fabales Malpighiales Oxalidales Celastrales Zygophyllales Saxifragales Vitales Dilleniaceae Solanales Gentianales Lamiales Boraginaceae Garryales Apiales Dipsacales Asterales Aquifoliales Ericales Cornales Caryophyllales Berberidopsidales Santalales Gunnerales Buxales Trochodendrales Proteales Sabiaceae Ranunculales Ceratophyllales Poales Commelinales Zingiberales Arecales Dasypogonaceae Asparagales Pandanales Dioscoreales Liliales Alismatales Acorales Magnoliales Laurales Piperales Canellales Chloranthales Austrobaileyales Nymphaeales Amborellales Pinales Cupressales Araucariales Welwitschiales Gnetales Ephedrales Cycadales Ginkgoales Selaginellales Isoetales Lycopodiales Polypodiales Cyatheales Marattiales Equisetales Psilotales Anthocerotophyta Bryophyta Marchantiophyta Zygnematophyceae Coleochaetophyceae Charophyceae Mesostigmatophyceae Chlorokyboph yceae Chlorophyceae Trebouxiophyceae p.p. Ulvophyceae p.p. Ulvophyceae p.p. Trebouxiophyceae p.p. Trebouxiophyceae p.p. Prasinophyceae p.p. Prasinophyceae p.p Figure 5 Fifty percent maximum likelihood majority-rule bootstrap consensus summary tree of Viridiplantae inferred from the all nucleotide positions (ntall) analysis. Data set derived from 78 protein-coding genes of the plastid genome (ntax = 360; 58,347 bp; missing data ~15.6%). Bootstrap support values 50% are indicated. Terminals with a triangle represent collapsed clades with > 2 taxa. Note position of Lycopodiophyta as sister to Spermatophyta is likely caused by base composition bias (see text). See Figures 9, 10, 11, 12, 13, and 14 for the complete tree and Additional file 1 for taxonomy. Lami. =Lamiidae; Campanuli. =Campanulidae; Lyco. =Lycopodiophyta. The placement of Dilleniaceae remains uncertain. The family is sister to Superrosidae in the ntall (95% BS), ntno3rd (77% BS), and RY (57% BS) analyses, but appears as sister to Superasteridae (70% BS) in the AA analysis. Within Superrosidae, a clade of Vitales +Saxifragales is supported in the ntall (75% BS), ntno3rd (70% BS), and AA (78% BS) analyses. In the RY analysis, the relationship among Saxifragales, Vitales, and remaining However, in the RY analysis, Proteales are sister to a clade containing Sabiaceae plus the remaining taxa, which has 79% BS. In the AA analysis, relationships among these three clades are unresolved. Among the remaining eudicots, we consistently recovered Trochodendrales as sister to Buxales+Pentapetalae Jari Oksanen (Ekologian yksikkö) 3 / 43 and Gunnerales as sister to the remaining lineages of Pentapetalae: Dilleniaceae, Superrosidae, andsuperasteridae. viherlevät Zygnematales Charales Coleochaetales maksasammalet Anthocerophyta 70,2 sammalet lieot saniaiset+kortteet 83,5 siemenkasvit 85,6 96,3 *,2 93,3 93,3 83,2 80,1 65,2 90,3 2,6 Sanikkaiset: hieman keinotekoinen nimi putkiloitiökasveille Pääjako: lieot vs. saniaiset + kortteet ja nämä lähempää sukua siemenkasveille Kulta-aika devoni - kivihiilikaudella nyt vain rippeet (etenkin Suomessa) Catharanthus roseus Allamanda cathartica Liekokasvit: Ipomoea purpurea lieot, mähkät ja Rosmarinus officinalis Tanacetum parthenium Inula helenium lahnaruohot Diospyros malabarica Kochia scoparia Carica papaya Eudicots Arabidopsis thaliana Hibiscus cannabinus Kortteet: Populus trichocarpa nykyään vain yksi suku Larrea tridentata Boehmeria nivea Medicago truncatula Vitis vinifera (Equisetum), Podophyllum peltatum kivihiilikaudella Aquilegia formosa Eschscholzia californica Sarcandra glabra kortepuita Chloranthales Liriodendron tulipifera Persea americana Houttuynia cordata Magnoliids Saruma henryi Zea mays : Sorghum bicolor suurin ja monimuotoisin Brachypodium distachyon Oryza sativa Sabal bermudana Monocots ryhmä Yucca filamentosa Smilax bona-nox Angiosperms (Flowering Plants) *,2 93,9 *,2 47,4 Colchicum autumnale Dioscorea villosa Jari Oksanen (Ekologian yksikkö) Acorus americanus 2 / 43 Kadsura heteroclite Nuphar advena Spermatophytes ANA Grade Amborella trichopoda Cunninghamia lanceolata (Seed Plants) Juniperus scopulorum Taxus baccata Sciadopitys verticillata Lieot ja *,11 sanikkaiset Yleiskuva Prumnopitys andina 89,10 Pinus taeda Cedrus libani Gymnosperms Gnetum montanum Euphyllophytes Welwitschia mirabilis Viherlevien siskoja Ephedra sinica Cycas micholitzii Cycas rumphii Tracheophytes Zamia vazquezii Ginkgo biloba (Vascular Plants) Psilotum nudum Ophioglossum petiolatum 87,8 Angiopteris evecta Monilophytes Pteridium aquilinum Cyathea spinulosa Equisetum diffusum 73,7 Dendrolycopodium obscurum Pseudolycopodiella caroliniana Huperzia squarrosa Lycophytes Embryophytes Selaginella moellendorffii (genome) Selaginella moellendorffii (1KP) (Land Plants) Leucodon sciuroides 81,12 Anomodon attenuatus 54,15 Rhynchostegium serrulatum Thuidium delicatulum 81,13 Bryum argenteum Rosulabryum capillare Mosses Hedwigia ciliata Ceratodon purpureus Streptophytes Physcomitrella patens Polytrichum commune 99,9 Sphagnum lescurii Marchantia polymorpha 80,6 Marchantia emarginata 97,8 Ricciocarpos natans Liverworts Sphaerocarpos texanus Bazzania trilobata Metzgeria crassipilis Nothoceros aenigmaticus Nothoceros vincentianus Hornworts Penium margaritaceum 62,5 98,2 Cosmarium ochthodes Roya obtusa Netrium digitus Spirogyra sp. Zygnematophyceae Cylindrocystis cushleckae 30,6 79,4 Mougeotia sp. Cylindrocystis brebissonii Mesotaenium endlicherianum 76,5 Coleochaete irregularis Coleochaete scutata Coleochaetales Chaetosphaeridium globosum Chara vulgaris Charales Entransia fimbriata Klebsormidium subtileklebsormidiales, Chlorokybales Spirotaenia minuta Chlorokybus atmophyticus Mesostigmatales Mesostigma viride Uronema sp. Nephroselmis pyriformis Pyramimonas parkeae Chlorophyta Monomastix opisthostigma Matrix type Supermatrix ASTRAL Fig. 3. Coalescent-based tree estimated from 424 gene trees estimated from first and second codon position alignments after removing genes with less Alignment AA DNA to AA DNA AA DNA to AA DNA 50% taxon occupancy after gene Codon fragments positions missing more NA than 66% 1 of and their 2 sites were all removed. NA Bootstrap NA 1 and 2 values all for the NA ASTRAL analysis are show Sister to land plants branches (* for %) along with the percentages of gene trees exhibiting significant conflict (bootstrap support >75%) with nodes discussed in the tex Zygnematophyceae-sister Charales-sister Figs. S8 and S9 for bootstrap values and Coleochaetales-sister percentages of conflicting gene trees for all nodes in analyses of first and second codon positions and amino respectively. Most nodes exhibited % Bryophytes bootstrap support in the ASTRAL analysis, but gray branches highlight nodes with less than % support. Mosses + liverworts models from sequenced genomes Wickett are Bryophytes etindicated al., monophyletic PNAS by E4859 E4868 blue text. (2014) blue: strong support, red: strong rejection Hornworts-sister Liverworts-basal Hornworts-basal Jari Oksanen (Ekologian yksikkö) 4 / 43 Gymnosperms

2 through the RNA-dependent DNA methylation pathway (10 12) andaccumulate Lieotmassively ja sanikkaiset in specific cells of the female gametophyte (13). Because the Selaginella srna population was generated from sporophytic tissues, the 24-nt sirna pathway may only be deployed during gametophyte development in Selaginella. A second distinction is the absence of DCL4, RDR6, Kladistiikkaa and MIR390 loci in Selaginella, which are required for the biogenesis of trans-acting sirnas (tasirnas) in angiosperms (2). Their absence suggests that tasirna-regulated processes in angiosperms, including leaf polarity (14) andde- velopmental phase changes in the sporophyte (15, 16), are regulated differently in Selaginella, and possibly reflects the independent origins of foliar organs in the lycophyte and angiosperm lineages (17, 18). Finally, the Selaginella plastome sequence reveals an extraordinarily large number of RNA-edited sites (2), as do other lycophyte organellar genomes (19, 20). This coincides with an exceptionally large number of PPR genes in Selaginella (>800) (2), some of which guide RNA editing events in angiosperms (21). Because Selaginella is a member of a vascular plant lineage that is sister to the euphyllophytes, we used comparative and phylogenetic approaches to identify gene origins and expansions coinciding with evolutionary innovations and losses in land plants. To identify such genes without regard to function, we compared the proteomes Lieot ja ofsanikkaiset the green alga Chlamydomonas, the moss Physcomitrella, Selaginella, and 15 angiosperm species; identified gene families that are related by homology by!"#"$%& new genes '()*+,"$"%-#& than the transition.%/0-".& from(12"& a plant3"2"1*"0& having a dominant 6-%(-$& gametophyte 21.#/*13& '*1$%.7& and simple,."'131%-$,& leaf- %("& *)#+8 1& 41.1* 0-#(+%+5)& less, and nonvascularized sporophyte (typified '()%".&9*"..&%(1$&:;&+<&"=%1$%&21.#/*13&'*1$%.>&<3+5&%(" by modern bryophytes) to a plant with a dominant, vascularized, and branched sporophyte "/'()**+'()%".&9?-,@&:A&!1/4".+$&B&C1$."$7&:DDEA&F"$8 3-#G&B&H31$"7&:DDIA&J3)"3&B&1*@7&EKK:17&EKKL17&4>@&M-28 with leaves. -$,&"/'()**+'()%".7&-$&%/3$7&#+5'3-."&%6+&51N+3&#*10".O In a second approach, we analyzed the phylogenies of genes known to function in Arabidopsis %("&.'"351%+'()%".&9.""0&'*1$%.>7&6(-#(&13"&-$&"=#"..&+< development (2). We identified 424 monophyletic EPK7KKK& groups.'"#-".& of developmental 9Q(+3$"7& genes, EKKEA& eachr#+%*1$0& group containing 1$0& putatively %("& 5+$-*+'()%".& all genes descended 9<"3$.7&."$./& from aj3)"3& B& 1*@7 B& S+3%*")7 EKKT>7& EKKL4>7& common 6-%(& land-plant 14+/%& D7KKK& ancestral.'"#-".7& gene (table -$#*/0-$,& S6). (+3."%1-*.7 6(-.G&<"3$.7&1$0&1**&"/.'+31$,-1%"&1$0&*"'%+.'+31$,-1%" Selaginella and Physcomitrella genes are present in 377 (89%) and 356 (84%) of the 424 land-plant <"3$.@&J*1$%.&%(1%&13"&-$#*/0"0&-$&%("&*)#+'()%"&1$0&<"3$ orthologous gene groups, respectively, indicating that the common ancestor of land plants had #*10".&13"&1**&.'+3"84"13-$,&+3&U.""08<3""V7&1$0&4"#1/." +<&%(-.&#+55+$&<"1%/3"&%("-3&5"54"3.&(12"&4""$&*/5'"0 most of the gene families known to direct angiosperm development. Conspicuous expansions %+,"%("3&(-.%+3-#1**)&/$0"3&213-+/.&%"35.7&./#(&1.&U'%"3-8 0+'()%".V& of families 1$0& within U<"3$.& different 1$0& lineages <"3$& 1**-".VW'131'()*"%-# resulted in different numbers of land-plant orthologs in each 1.."54*1,".&+<&'*1$%.@&Q("&<+#/.&+<&%(-.&3"#*1..-<-#1%-+$ genome (table S6). The 27 vascular plant-specific -.& "=#*/.-2"*)& orthologous groups +$& <"3$.& likely9x-2-.-+$& represent genes Q31#("+'()%17& associatedy/'()**+'()%-$17& with developmental innovations Z$< $& of vascu- [+$-*-<+35+'8 R/ $&.".7&+<&F"$3-#G&B&H31$"7&:DDI>7&#(131#%"3-\"0&4)&*1%"31* lar plants. Among them are genes regulating the 3++%& Yleiskuva meristem (CLV1 and CLV2), hormone signaling +3-,-$& -$& %("& "$0+0"35-.7& /./1**)& 5".13#(& '3+8 (GID1 and CTR1), and flowering (TFL2 and %+=)*"5&-$&.(++%.7&1&'."/0+"$0+.'+3"7&'* *&%1'"8 UFO). Homologs of genes involved in the specification of xylem (NST and VND) (22) and %/57&1$0&.'"35&#"**.&6-%(&TK]:KKK&<*1,"**1"&9!"$\1,*-1 B&1*@7&EKKKA&R#($"-0"3&B&1*@7&EKKE1>@ Z$#3"1.-$,*)& 3+4/.%& '()*+,"$"%-#& ()'+%(".".& <+3 Jari Oksanen (Ekologian yksikkö) 5 / 43 Maakasvien evoluutio A liverworts mosses Physcomitrella hornworts bryophytes clubmosses lycophytes non-vascular land plants quillworts spikemosses Selaginella ferns gymnosperms euphyllophytes vascular plants of far fewer new genes (516) than the transition (AS2 and ULT1), hormone signaling and biosynthesis (BRI1, BSU1, ARF16, ACS, and ACO), Yleiskuva -.,/%$&01 Yleiskuva from a basal vascular #*1..-<-#1%-+$7&H-4+%-1#"1"7&<"3$.7&5+$-*+'()%".7&5+$+'()*"%-#@ plant to a basal euphyllophyte whose descendants include the angiosperms (1350). and flowering (HUA1, EMF1, FT, TFL1, and These numbers show that the evolution of traits FD). Altogether, these Putkilokasvit: results suggest that konsensus the specific!"#$%&'(#!%" to euphyllophytes or angiosperms required the evolution of about three times more to a vascular angiosperm included the stepwise Euphyllophytes evolutionary transitions from a nonvascular plant addition of components of some developmental pathways, especially those Lycophytes regulating meristem Spermatophytes Monilophytes and hormone biology, as previously noted for the gibberellin signaling pathway (24, 25). Genes involved in secondary metabolism were also investigated because plants synthesize numerous secondary metabolites that they use to interact with their environment. Three gene families involved in their biosynthesis, including those encoding cytochrome P450-dependent monooxygenases (P450s), BAHD acyltransferases (BAHDs), and terpene synthases (TSs), were analyzed. The largest of these in Selaginella is the P450 family, accounting for 1% of its predicted proteome (table!"#$% &$% S7) '()*+)*,*% (2). All three families -./0(#+)/% Smith 1+-"23")#% AR et al., Taxon 4+053"()*."-*% 55, ; 2006 (6 show similar evolutionary 758(4%95*2,054%-05)3%0")+5#+*$%:(-(0(#/%*,7754";+*%3.+ trends, with the inferred ancestral 4+*,03*% vascular plant (6% -4+9"(,*0/% having a small -,<0"*.+1% -./0(#+)+3"2% *3,1"+* number of genes =+$#$>%?5,<+*()%@%A5)*+)>%&BBCD%E+)4"2F%@%'45)+>%&BBGD that radiated extensively but independently within the lycophyte and angiosperm lineages (figs. S6 to S13). BAHD and TS?+);5#0"5%@%50$>%CHHHD%I4/+4%@%50$>%CHH&5>%*++%75")%3+J3%6(4 (3.+4*K$%?+*(09+1%)(1+*%.59+%4+2+"9+1%<((3*345-%*,--(43 genes, which are known to be involved in the GH$ biosynthesis of volatile odorants, are apparent only in seed plants (figs. S12 to S13), likely reflecting the coevolution <"3$.&9^1."4"&B&1*@7&:DDL7&:DD_A&[1$(13%7&:DDL7&:DD_A of seed plants with animals that pollinate flowers or disperse seeds. The J3)"3&B&1*@7&:DD_7&EKK:17&EKKL4A&F31$\&B&^/..7&:DDPA Yleiskuva independent diversification of these gene families plus the large J1($G"& number B& of Selaginella 1*@7& :DDPA& genes S+*<7& :DDIA& S+*<& B& 1*@7& :DD`A suggest that Selaginella a"#g"3%&b&1*@7&:ddda&b1$,"3+6&b&1*@7&:ddda&r1$+&b&1*@7 not only has the potential to synthesize EKKK1A&R#($"-0"3&B&1*@7&EKKL#A&S-G.%3c5&B&J3)"37&EKK_A a repertoire of secondary metabolites that rivals Q./%./5-&B&F1%+7&EKKPA&R#(/"%%'"*\&B&1*@7&-$&'3"..>7&/%-8 the angiosperms in complexity, but that many of them are likely to be B unique. Some have been shown to be of pharmaceutical value [e.g., (26)]. IK_ 15 Angiosperms Monocots Eudicots Selaginella We have used the compact Selaginella genome sequence to uncover genes associated with Physcomitrella 8155 Basal angiosperm gene set major evolutionary transitions in land plants. Understanding their functions in Selaginella and other taxa, as well as acquiring the genome sequences of other informative taxa, especially 7247 Basal vascular plant gene set Chlamydomonas charophytes, ferns, and gymnosperms, will be key 6820 Basal land plant gene set to understanding the evolution of plant form and function. angiosperms 15 species Liekokasvit ovat erillinen ryhmä + kortteet ( Monilophyta ) ovat siemenkasvien sisarryhmä Silmiinpistävin ero on eufyllofyyttien varsinainen lehti vs. liekojen mikrofylli Putkilokasvien ( trakeofyyttien ) yleisiä piirteitä Elinkierto heteromorfinen, itsenäinen sporofyytti näkyvin osa Alkionkehitys useimmiten suojassa (kehittyneemmillä ryhmillä) Yleensä heterosporia: megasporit ja mikrosporit eli isoitiöt ja pikkuitiöt Jari Oksanen (Ekologian yksikkö) 6 / 43 chlorophytes charophytes 3814 Basal green plant gene set Chlamydomonas References and Notes 1. P. Kenrick, P. R. Crane, Nature 389, 33 (1997). Fig. 2. (A) Phylogeny ofplants. Taxainred have sequenced genomes. (B) Genefamily gains(+) and 2. Details are given in the supporting Fig. 1. materials The on green Sciencelineage and the AM symbiosis. (A) Schematic representation of the green lineage. Light blue: chlorophytes and basal charophytes; losses ( ) mappedontotheplantphylogenetictree.theminimumnumbersofgenefamiliespresentin Online. the ancestors of different plant lineages are circled. 3. W. Wang et al., BMC Plant Biol. 5, 10 (2005). dark blue: advanced charophytes. Green bars on the left indicate the actual Banks muuta, Science 332, (2011) number of transcriptomes and genomes included in the extended dataset MAY 2011 VOL 332 SCIENCE used in this study. (B) Schematic representation of the symbiotic genes. Jari Oksanen (Ekologian yksikkö) 7 / 43 Symbioosi AM-sienten kanssa edellytys maalle siirtymiseen Delaux et al, PNAS 112, (2015) Jari Oksanen (Ekologian yksikkö) 8 / 43 Results Extensive Genes in tion from sets avai at this tr ning the dition to dataset a liverw complex Closteriu (Spirogyr mosses, a project tended d starting RLK, DM

3 Liekokasvit: nykyiset lajit Liekokasvit Nykyiset lajit Liekokasvit: Pääryhmät Liekokasvit Nykyiset lajit Sporangiot lateraaliset, mikrofyllien pinnalla Sekä homosporisia että heterosporisia Kehittynyt heteromorfia: gametofyytti jää pieneksi, usein itiöiden sisällä Selaginella: gametofyytti Lycopodiopsida Selaginellidae Lycopodidae Selaginellales Isoëtales Jari Oksanen (Ekologian yksikkö) 9 / 43 Yksi [tai kaksi] alaluokka[a] Lieot ([Lycopodidae:] Lycopodiales) Yksi heimo (Lycopodiaceae), 1 3 sukua 1 (Huperzia, Lycopodiella, Lycopodium) kasvioissa ehkä ylipilkottu Homosporisia Korkea kromosomiluku (jopa 2n = 134) muinaisten liekokasvien allopolyploideja? Ei paksuuskasvua Mähkät ([Selaginellidae:] Selaginellales) ainoa suku Selaginella Mikrofyllissä kieleke Heterosporia Lahnaruohot ([Selaginellidae:] Isoëtaceae) ainoa suku Isoëtes Lyhyt maavarsi, jossa paksuukasvu ytimestä: uloimmat osat vanhimmat Lehdet mikrofylleiksi isot, kielekkeelliset Heterosporia 1 Christenhusz et al., Phytotaxa 19, 17 54; 2011 Jari Oksanen (Ekologian yksikkö) 10 / 43 Liekokasvit Sukupuuttoon kuolleet ryhmät Fossiiliset ryhmät yleistä Leclerqia (keskidevoni) Pleuromeia (trias) Varhaisimmat maakasvit matalia ruohoja (Zosterophyllum) Vanhemmat muodot (Protolepidodendrales: Leclercqia complexa) muistuttivat nykyisiä liekoja, homosporisia Suuria (40 m) liekopuita: Lepidodendron (Lepidodendrales) Strobilus ( käpy ), jossa iso- ja pikkutiölehtiä Lepidocarpon (Lepidodendrales): mikrofylli kasvoi megasporangion suojaksi siemenmäinen Lepidophloios: iso- ja pikkuitiöstrobilukset erilaiset kaksikotinen Jari Oksanen (Ekologian yksikkö) 15 / 43 Polypodiopsida Psilotidae Polypodiidae Equisitidae Marattiidae Ophioglossales Psilotales Kortteita pidettiin ennen erillisenä pääryhmänä, mutta muodostavat yhdessä saniaisten kanssa monofyleettisen ryhmän, joka on siemenkasvien sisar Ennen primitiivisimpänä elävänä kasvina pidetty Psilotum (haarusanikkainen) näyttääkin olevan läheinen käärmeenkielille (Ophioglossum) ja noidanlukoille (Botrychium) mutta tässä kohtaa analyysi on epävarma Suurin nykyinen ryhmä on köykäispesäkkeiset saniaiset (Polypodiidae) Toiseen saniaisryhmään (Marattidae) kuuluu lähinnä trooppisia lajeja Jari Oksanen (Ekologian yksikkö) 21 / 43

4 letters to nature Sanikkaispuu yleistä We obtained DNA sequences (5,072 aligned base pairs) of four genes from two plant genomes: plastid atpb, rbcl and rps4, and nuclear small-subunit ribosomal DNA. We also assembled a congruent data set of 136 vegetative and reproductive morphological/ anatomical characters. We sampled 35 representatives from all major monophyletic lineages of land plants. The selection of taxa re ects our focus on basal vascular plants, and all six Euphyllophytina 1 be very useful in inferring deep phylogenetic patterns 4±6. With few exceptions 12,20, broad phylogenetic studies rely solely on combined nucleotide sequence data, with authors arguing that morphological character homology assessment among ancient and divergent groups is too challenging. This practice ignores the higher complexity of morphological characters that can conserve character states over time and that have a lower probability of random evolution of Pryer et al., Nature 409, ; 2001 similar structures. lineages are represented by two or more members. Five bryophytes Megafylli yleistä /96 /98 75/93 /99 94/59 87/56 / / / / / / / / -/64 Isoetes Huperzia 70/81 92/ Polytrichum Sphagnum 88/96 Cycas Ginkgo Anthoceros / Osmunda / Angiopteris Marattia Danaea Psilotum Tmesipteris Pinus 90/91 62/- 55/- 89/- 65/- 92/- 91/- 69/- -/- 86/ / Dicksonia Plagiogyria Cyathea Phanerosorus Gleichenia Hymenophyllum Ophioglossum Botrychium Austrobaileya Chloranthus Marchantia / Osmunda Marattiidae Haplomitrium Pteridium Blechnum Marsilea Lygodium NATURE VOL FEBRUARY Macmillan Magazines Ltd 619 tree ferns Salvinia Equisetum 1 Equisetum 2 Ophioglossidae Psilotidae whisk ferns Spermatophytata seed plants Selaginella bryophytes gleichenioid ferns eusporangiate ferns Gnetum polypodiaceous ferns Equisetopsida horsetails Lycophytina heterosporous water ferns Spermatophytata Lycophytina Bryophytes Polypodiidae leptosporangiate ferns Moniliformopses 0.1 substitutions per site Jari Oksanen (Ekologian yksikkö) 22 / 43 Figure 1 Phylogenetic relationships for all the main lineages of vascular plants inferred them are arbitrary. Branches leading to the three monophyletic clades of vascular plants from maximum-likelihood (ML)analysis of the combined chloroplast rbcl, atpb, rps4 and (lycophytes, seed plants and horsetails+ferns) are drawn medium thick. The branch nuclear small-subunit rdna data set. Numbers at nodes and before the slash are ML supporting the Euphyllophytina, with horsetails+ferns as sister group to seed plants, is the bootstrap values $50%; maximum parsimony (MP) bootstrap values $50% appear after thickest. Wiggled lines (at straight arrows) indicate three areas of con ict between the ML the slash when these same nodes were supported in the MP unequally weighted analysis and MP analyses. Branch lengths are proportional ja kortteet to number of substitutions yleistä per site of the combined four-genes plus morphology data set (single MP tree = steps). (scale bar). Thumbnail sketches of plant representatives accompany major clades. A minus sign indicates a node had less than 50% bootstrap support in one or the other Taxonomy follows ref. 1. analysis. The topology is rooted by all bryophytes, hence relationships depicted among Megafylli ja steeli Euphyllophytina Polypodiidae: köykäispesäkkeiset sanikkaiset, joilla itiöpesäke syntyy yhdestä solusta Moniliiformes: saniaiset ja kortteet yhdistävä ryhmä Euphyllophytina: kasveilla varsinainen lehti eli megafylli Spermatophyta: siemenkasvit Lycophytina: liekokasvit Bryophytes: sammalet ja maksasammalet Liekokasvien lehti on mikrofylli, jolla ei ole yhteyttä keskuslieriöön Saniaisten lehden johtosolukko on kehittynyt ja sen kanta tekee aukon keskuslieriöön Telomiteorian mukaan lehti on syntynyt litistyneestä ja yhteenkasvaneesta versosta Kortteidein lehdet näyttävät mikrofylleiltä, sillä niissä on vain yksi keskisuoni, mutta fossiiliaineistojen mukaan ne ovat kehittyneet megafylleistä Jari Oksanen (Ekologian yksikkö) 23 / 43 Käärmeenkielikasvit (Ophioglossales) Käärmeenkielet ja haarusanikkaiset Viisi sukua, joista Suomessa käärmeenkieli (Ophioglossum vulgatum) ja noidanlukkoja (Botrychium) Lehti ei tasossa, vaan fertiili ja steriili osa erikseen Maavaressa sekundaarista paksuuskasvua Jopa yli 0 kromosomia muinaisia allopolyploideja? Ei fossiiliaineistoa Asema ja suhde haarusanikkaisiin vielä epävarma Jari Oksanen (Ekologian yksikkö) 24 / 43 Jari Oksanen (Ekologian yksikkö) 25 / 43

5 Haarusanikkaiset (Psilotales) Käärmeenkielet ja haarusanikkaiset Kortteet (Equisetopsida) Kortekasvit Psilotum: dikotomisesti haarautunut yhteyttävä varsi Mikrofyllimäiset lehdet Pidetty (ennen) kaikkein primitiivisimpänä elävänä sanikkaisena: muistuttaa ulkoisesti ensimmäisiä maakasveja (Rhynia) Kolmilokeroiset, kehittyneet sporangiot eivät sopineet kuvaan Sekvenssiaineisto yhdistää saniaisiin: ilmeisesti yksinkertaistunut Yhdessä toisen suvun (Tmesipteris) kanssa trooppisia epifyyttejä Psilotum nudum sporangiot (OY puutarha) Nykyään vain Equisetum-suku (n. 32 lajia) Päätteinen kasvu, mikrofyllimäiset lehdet, strobilus latvassa Homosporisia, mutta itiöistä kahdentyyppisiä yhteyttäviä gametofyyttejä: Suurin osa jää hyvin pieniksi ja tuottaa vain anteridioita Muutamat jatkavat kasvuaan ja tuottavat arkegonioita jopa pari vuotta Jos ei hedelmöitystä, voivat lopulta tuottaa myös anteridioita Fossiilisilla muodolla myös heterosporiaa Jari Oksanen (Ekologian yksikkö) 26 / 43 Jari Oksanen (Ekologian yksikkö) 28 / 43 Kortekasvit Korte (Equisetum): kasvullinen rakenne Kortekasvit: Fossiilisia muotoja Kortekasvit Useita sukupuuttoon kuolleita ryhmiä, joukossa kookkaita puita Trimerophytopsida: monopodiaalinen varsi, lienee kehittynyt dikotomisesta telomista Cladoxylopsida (Calamophyton): lyhytvartisia puita ja liuskoittunut steeli Calamitales: puita, joilla kortemainen strobilus Calamophyton (devoni) Jari Oksanen (Ekologian yksikkö) 29 / 43 Jari Oksanen (Ekologian yksikkö) 31 / 43

6 Marattiidae Ko yka ispesa kkeiset saniaiset Ainakin lajia ja > 40 heimoa2, myo s enimma t suomalaiset lajit Ko yka ispesa kkeinen (leptosporangiaattinen): itio pesa ke syntyy yhdesta solusta Itio pesa kkeet yleensa ryhmissa (sorus), katesuomun (indusium) suojaamina Useimmat homosporisia, gametofyytti lehtima inen Pieni trooppinen ryhma (mutta kookkaita, na ytta via lajeja) Itio pesa ke syntyy monesta solusta: eusporangiaattinen Fossiileja ainakin hiilikaudelta Marattia 2 33 / 43 Christenhusz et al. Phytotaxa 19, 7 54; 2011 Ko yka ispesa kkeiset saniaiset kehittyiva t kukkakasvien varjossa 35 / 43 Modernien saniaisten kehitys letters to nature Schneider et al. Nature 428, (2004) Ko yka ispesa kkeisten saniaisten pa a ryhma t kehittyiva t samaan aikaan kuin koppisiemeniset: moderneja ryhmia Schneider et al. Nature 428, (2004): Lineages through time 38 / / 43

7 Vesisaniaiset (Salviniales): heterosporia Jari Oksanen (Ekologian yksikkö) 40 / 43