Tracheophyte Phylogeny Poster Vascular – Systematics and Characteristics

Hornworts, Mosses, Liverworts see "Bryophyte" Phylogeny Poster

homosporous lvs eligulate sporangia s.t. clustered in terminal strobili Lycopodiales Lycopodiaceae lvs usually with single central, unbranched vein root dichotomous cormose to rhizomatous sporangia reniform, dorsiventral and transversely dehiscent, 1 (adaxial) per 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

sporophyte dominant receptacle elongate sporangia vascular tissue: sporangia in sori lvs thin, usually 1 cell layer single-layered tracheids + sieve cells sporangia primarily sori marginal with annulus: Hymenophyllaceae sporangia many multi-layered: eusporangial Hymenophyllales zygote: tapetum plasmodial leptosporangial 1st division horizontal pseudoendospore + siphonostele root steles with 3–5 protoxylem poles stem protoxylem mesarch 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 ferns Liverworts Cibotiaceae Dicksoniaceae Metaxyaceae sori often terminal on veins Cyatheales Mosses Thyrsopteridaceae Loxsomataceae

Hornworts Culcitaceae Plagiogyriaceae

Lycophytes

Ferns Saccolomataceae Cystodiaceae Tracheophytes (incl. horsetails) Cycads sporangial maturation mixed Lonchitidaceae Lindsaeaceae Ginkgo annulus vertical on sporangium euphyll Pteridaceae Dennstaedtiaceae Ephedra interrupted by stalk and stomium leaf gaps Welwitschia Polypodiales protoxylem exarch Gnetum Gymnosperms Eupolypods I: Polypodiineae roots monopodial Plants (incl. Polypodiaceae, Davalliaceae, variously vessels ANA grade 30-kb cp inversion Didymochlaenaceae, Dryopteridaceae, Hypodematiaceae, Angiosperms Magnoliids Lomariopsidaceae, Oleandraceae, Tectariaceae) Monocots

Fabids Eupolypods II: Aspleniineae

alvids (incl. Aspleniaceae, Athyriaceae, Blechnaceae, Rosids M Cystopteridaceae, Desmophlebiaceae, Diplaziopsidaceae, Lamiids Hemidictyaceae, Onocleaceae, Rhachidosoraceae, Campanulids Asterids Thelypteridaceae, Woodsiaceae)

Euphyllophytes males: whorls of megasporophylls

at trunk apex alternate with trophophylls (no seed cones) Cycadaceae Cycas pachycaulous; pinnate megaphylls males with pollen cones; pollen tube simple roots with N -fixing cyanobacteria 2 seed cones with 2[3] per megasporophyll Bowenia Ceratozamia Chigua Dioon lignins with syringaldehydes Cycadales Zamiaceae Encephalartos Lepidozamia Macrozamia dioecious Microcycas Stangeria Zamia stout short 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 ovules/seeds not Ephedraceae Ephedra enclosed by carpel pollen tube haustorial vessels porose caudex primarily 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 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 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 Seed cotyledons 2 Podocarpaceae Microcachrys Nageia Parasitaxus Pherosphaera Phyllocladus Plants Podocarpus Prumnopitys Retrophyllum Cupressales monoecious eustele "double needles" (cladodes) secondary growth ovules 7–9 Sciadopityaceae Sciadopitys heterosporous Conifers II ovules / seeds dioecious pollen evergreen; pollen cones tiny pollen tube peltate microsporangiophores with 2 gametes each 1 ovule, 1 seed/cone, arillate Amentotaxus Austrotaxus Cephalotaxus cotyledons 2 Taxaceae Pseudotaxus Taxus Torreya

monoecious cone scales (Juniperus: monoecious or dioecious) opposite lvs mostly scale-like pollen not saccate ovules 1–20/scale cotyledons many seed cones terminal Juniperus Neocallitropsis

Spermatophytes Pilgerodendron 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 (2018) Tracheophyte Phylogeny Poster • hypothetical tree based on molecular phylogenetic data (Oct. 2018) • 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 • this poster is now available in 12 world languages

CC-BY Dipl. Biol. Theodor C. H. Cole ( Prof. Dr. Julien B. Bachelier References Angiosperm Tracheophyte Bryophyte Schuettpelz, Schneider, The Pteridophyte Phylogeny Group, 2016 Prof. Dr. Hartmut H. Hilger 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 Sciences (DCPS) Phylogeny Phylogeny Phylogeny Stevens 2018, APweb, www.mobot.org/MOBOT/research/APweb Institute of Biology – Botany Textbooks: Judd et al. 2016, Simpson 2010

Freie Universität Berlin Poster Poster Poster Thanks to all members of the TPP team: Fernanda Antunes Carvalho, Mohamed Fennane, Diego Medan, Sergei L. Mosyakin, Sofi Mursidawati, Dashzeveg Nyambayar, Anastasiya V. Odintsova, Batlai Oyuntsetseg, Federico Selvi, Dmitry D. Sokoloff, Xin Zhong

© The Authors, 2018 © The Altensteinstr. 6, D-14195 Berlin, Germany