Tracheophyte Phylogeny Poster VASCULAR PLANTS – SYSTEMATICS AND CHARACTERISTICS HORNWORTS, MOSSES, LIVERWORTS SEE "BRYOPHYTE" PHYLOGENY POSTER homosporous lvs eligulate sporangia sometimes clustered in terminal strobili LYCOPODIALES Lycopodiaceae lvs usu. with single central, unbranched vein root dichotomous cormose to rhizomatous sporangia reniform, dorsiventral and transversely dehiscent, 1 (adaxial) per leaf lvs spiral, in basal rosette microspores monolete megaspores 50–300 ISOËTALES Isoëtaceae heterosporous lvs ligulate sporangia on 4-sided strobili lvs 4-ranked microspores trilete megaspores 4 SELAGINELLALES Selaginellaceae sporangiophores in terminal strobili lvs whorled, fused into sheaths encircling base of internodes stems ridged with internal hollow canals spores with elaters 4–6, straplike EQUISETALES Equisetaceae Lycophytes each leaf with fertile (sporophore) & sterile (trophophore) parts (latter inclined relative to former) Ophioglossaceae roots unbranched, root hairs absent OPHIOGLOSSALES Ferns collateral leaf vascular bundles gametophyte nonphotosynthetic, often subterranean, mycorrhizal roots absent lvs reduced, veins 1 or 0 sporangia 2–3, fused: synangium PSILOTALES Psilotaceae tropical, megaphyllous, stipules + polycyclic siphonostele often synangia, annulus – sporangia open by longitudinal slit or pores MARATTIALES Marattiaceae annulus lateral on sporangium opening apically OSMUNDALES Osmundaceae Monilophytes receptacle elongate sporangia sporangia in sori lvs thin, usually 1 cell layer sporophyte dominant single-layered sporangia primarily multi-layered: sori marginal vascular tissue: with annulus: YMENOPHYLLALES Hymenophyllaceae tracheids + sieve cells eusporangial tapetum plasmodial H sporangia many pseudoendospore + leptosporangial zygote: siphonostele – 1st division horizontal stem protoxylem mesarch root steles with 3 5 protoxylem poles rhizome with scales; veins anastomosing sporangia maturation simultaneous GLEICHENIALES Gleicheniaceae Matoniaceae Dipteridaceae annulus transverse, subapical opening by longitudinal slit SCHIZAEALES Anemiaceae Lygodiaceae Schizaeaceae aquatic aerenchyma + leaf dimorphism heterosporous; sporocarps; annulus – hairs SALVINIALES Marsileaceae Salviniaceae endospore 2-layered Cyatheaceae usually tree ferns Cibotiaceae Dicksoniaceae Metaxyaceae sori often terminal on veins CYATHEALES Thyrsopteridaceae Loxsomataceae Culcitaceae Plagiogyriaceae LIVERWORTS MOSSES Saccolomataceae Cystodiaceae HORNWORTS sporangial maturation mixed Lonchitidaceae Lindsaeaceae LYCOPHYTES annulus vertical on sporangium Pteridaceae Dennstaedtiaceae FERNS interrupted by stalk and stomium RACHEOPHYTES (incl. HORSETAILS) OLYPODIALES T P Eupolypods I: Polypodiineae CYCADS GINKGO (incl. Polypodiaceae, Davalliaceae, EPHEDRA WELWITSCHIA Didymochlaenaceae, Dryopteridaceae, Hypodematiaceae, GNETUM EED GYMNOSPERMS S CONIFERS Lomariopsidaceae, Oleandraceae, Tectariaceae) PLANTS euphyll ANA GRADE leaf gaps Eupolypods II: Aspleniineae ANGIOSPERMS MONOCOTS protoxylem exarch (incl. Aspleniaceae, Athyriaceae, Blechnaceae, roots monopodial MAGNOLIIDS variously vessels Cystopteridaceae, Desmophlebiaceae, Diplaziopsidaceae, 30-kb cp inversion FABIDS Hemidictyaceae, Onocleaceae, Rhachidosoraceae, ALVIDS ROSIDS M Thelypteridaceae, Woodsiaceae) LAMIIDS AMPANULIDS ASTERIDS C females: whorls of megasporophylls Euphyllophytes at trunk apex alternate with trophophylls (no seed cones) Cycadaceae Cycas pachycaulous; pinnate megaphylls males with pollen cones; pollen tube simple roots with N2-fixing cyanobacteria seed cones with 2[3] seeds per megasporophyll Bowenia Ceratozamia Chigua Dioon lignins with syringaldehydes CYCADALES Zamiaceae Encephalartos Lepidozamia Macrozamia dioecious Microcycas Stangeria Zamia stout short shoots with zooidogamy lvs flabelliform, dichotomously veined, deciduous INKGOALES pollen tube branched G ovules 2 (basal collar); cotyledons 2 Ginkgoaceae Ginkgo xeromorphic, lvs scale-like, sheathed, shed early stems narrow, striate, photosynthetic seed cones: 1–3 ovules; double fertilization Ephedra ovules/seeds not Ephedraceae enclosed by carpel pollen tube haustorial vessels caudex primarily porose perforation plates lvs 2 (straplike) life-long basal continuous growth anemophilous dioecious venation parallel primary endosperm ♀ gametophyte tubes grow towards pollen tubes Welwitschia no bisexual strobili striate pollen Welwitschiaceae binucleate sperm GNETALES cotyledons 2 ectomycorrhizal lvs opposite, simple, broad, venation reticulate (angiosperm-like "convergence") laticifers Gnetaceae Gnetum monoecious resin canals; lvs linear (needles) pollen mostly 2-saccate Abies Cathaya Cedrus Hesperopeuce PINALES ovules 2, inverted seeds winged Pinaceae Keteleeria Larix Nothotsuga Picea loss of Pinus Pseudolarix Pseudotsuga Tsuga spermatozoids monoecious or dioecious Conifers I lvs broad to acicular nucellar pollen not saccate siphonogamy seed cones large, disintegrate cotyledons 2–4 Agathis Araucaria Wollemia leaves Araucariaceae Gymnosperms morphologically dioecious; roots with nodules diverse 1 ovule/scale, cone often one seeded receptacle and/or Acmopyle Afrocarpus Dacrycarpus Dacrydium Falcatifolium epimatium/carpidium fleshy Halocarpus Lagarostrobus Lepidothamnus Manoao cotyledons 2 Seed Podocarpaceae Microcachrys Nageia Parasitaxus Pherosphaera Phyllocladus Plants Podocarpus Prumnopitys Retrophyllum Saxegothaea CUPRESSALES monoecious "double needles" (cladodes) eustele ovules 7–9 Sciadopityaceae Sciadopitys secondary growth Conifers II heterosporous dioecious ovules evergreen; pollen cones tiny seeds peltate microsporangiophores pollen 1 ovule, 1 seed/cone, arillate Amentotaxus Austrotaxus Cephalotaxus pollen tube with 2 gametes each cotyledons 2 Taxaceae Pseudotaxus Taxus Torreya monoecious cone scales (Juniperus: monoecious or dioecious) opposite lvs mostly scale-like Actinostrobus Athrotaxis Austrocedrus Callitris pollen not saccate Calocedrus Chamaecyparis Cryptomeria Cunninghamia ovules 1–20/scale cotyledons many Cupressus Diselma Glyptostrobus Fitzroya Fokienia seed cones terminal Cupressaceae Juniperus Libocedrus Metasequoia Neocallitropsis Spermatophytes Papuacedrus Pilgerodendron Platycladus Sequoia Sequoiadendron Taiwania Taxodium Tetraclinis Thuja Thujopsis Widdringtonia Xanthocyparis bisexual flower; fruit (ovules enclosed by carpel); loss of spermatozoids pollen tube penetrating stigma/style/nucellus/synergid (complete siphonogamy) vessels; pollenkitt; primarily zoophilous double fertilization: triploid endosperm ANGIOSPERMS SEE ANGIOSPERM PHYLOGENY POSTER COLE TCH, BACHELIER JB, HILGER HH (2020) Tracheophyte Phylogeny Poster • hypothetical tree based on molecular phylogenetic data (2020) • branch lengths deliberate, not expressing actual time scale • if a character is marked as being a potential synapomorphy at a node/for a clade, ) this does not mean that all members of that clade possess that character Dipl. Biol. Theodor C. H. Cole • this poster is now available in 20 languages CC-BY Prof. Dr. Julien B. Bachelier References ( Schuettpelz, Schneider, The Pteridophyte Phylogeny Group, 2016 Prof. Dr. Hartmut H. Hilger Angiosperm Tracheophyte Bryophyte Kubitzki (ed.) 1990, FGVP, Vol. I, Pteridophytes and Gymnosperms Christenhusz et al. 2011/2014; Soltis et al. 2005/2011; Ran et al. 2018 Dahlem Centre of Plant Sciences (DCPS) Stevens 2019: APweb, www.mobot.org/MOBOT/research/APweb Institute of Biology – Botany Phylogeny Phylogeny Phylogeny Textbooks: Judd et al. 2016, Simpson 2020 Freie Universität Berlin Thanks to all members of the TPP team: Fernanda Antunes Carvalho, Johannes Enroth, Mohamed Fennane, Milan Gavrilović, Bui Thu Ha, Poster Poster Poster Michal Hroneš, Peđa Janaćković, Joo-Hwan Kim, Ľuboš Majeský, Pavol Mártonfi, Diego Medan, Sergei L. Mosyakin, Sofi Mursidawati, Sławomir Nowak, Altensteinstr. 6 Dashzeveg Nyambayar, Anastasiya V. Odintsova, Batlai Oyuntsetseg, Federico Selvi, Gabriella Somogyi, Magsar Urgamal, Xin Zhong © The Authors, 2020 © The D-14195 Berlin, Germany.