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Tracheophyte Phylogeny Poster (TPP) -

Vascular Plants: Systematics and
Characteristics, 2016

Data · July 2016

DOI: 10.13140/RG.2.1.1947.5443

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2 authors:

Theodor C. H. Cole Hartmut H. Hilger

Freie Universität Berlin Freie Universität Berlin
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 Tracheophyte Phylogeny

Vascular Plants – Systematics and Characteristics

H ornworts, Mosses, Liverworts see "B ryophyte " P hylogeny P oster

homosporous

Lycopodiales
lvs eligulate
sporangia reniform, basal on sporophyll or in terminal strobili Lycopodiaceae
shoots with lycophylls
dichopodial root: protoxylem endarch cormose to rhizotamous
stem protoxylem exarch lvs spiral, in basal rosette

Isoëtales
sporangia dorsiventral and transversely dehiscent microspores monolete
megaspores 50–300 Isoëtaceae
heterosporous
sporangia in axils of sporophylls
Lycophytes

lvs ligulate
on 4-sided strobili; lvs 4-ranked

Selaginellales
microspores trilete
megaspores 4 Selaginellaceae
sporangiophores in terminal strobili
lvs whorled, fused into sheaths at base

Equisetales
stems ridged with internal hollow canals
spores with elaters 4–6, straplike Equisetaceae

Ophioglossales
sporophores: each leaf with sterile & fertile segments
(latter inclined relative to former)
Ophioglossaceae
Ferns roots unbranched, root hairs absent
collateral leaf vascular bundles
gametophyte nonphotosynthetic, often subterranean, mycorrhizal
roots absent

Psilotales
lvs reduced, veins 1 or 0
sporangia 2–3, fused: synangium Psilotaceae
Monilophytes

polycyclic siphonostele
Marattiales Marattiaceae

annulus on one side of sporangium

Osmundales Osmundaceae

sporophyte dominant
vascular tissue: receptacle elongate

Hymenophyllales
tracheids + sieve cells lvs thin, usually 1 cell layer
sporangia many sporangia in sori annulus sori marginal Hymenophyllaceae
zygote: tapetum plasmodial
1. division horizontal pseudoendospore + leptosporangia
siphonostele
variously vessels root steles with 3–5 protoxylem poles

Gleicheniales
stem protoxylem mesarch rhizome with scales; veins anastomosing
sporangia maturation simultaneous Gleicheniaceae

annulus transverse, subapical

Schizaeales Anemiaceae Lygodiaceae Schizaeaceae

aquatic; aerenchyma +

Salviniales
stems dichotomizing; leaf dimorphism
hairs heterosporous; sporocarps; annulus – Marsileaceae Salviniaceae
endospore
2-layered

Cyatheales
hairs +
sori terminal on veins Cyatheaceae

Liverworts

Mosses
Eupolypods I: Polypodiaceae*
sporangial maturation mixed (incl. Polypodioideae, Davallioideae,
Hornworts

Polypodiales Didymochlaenoideae, Dryopteridoideae, Hypodematioideae,

sporangium with vertical annulus
euphyll interrupted by stalk and stomium
leaf gaps
Lycophytes Lomariopsidoideae, Oleandroideae, Tectarioideae)
protoxylem exarch Ferns
roots monopodial
(incl. horsetails)
Eupolypods II: Aspleniaceae*
30-kb cp inversion Palmferns
Ginkgo (incl. Asplenioideae, Athyrioideae, Blechnoideae,
Ephedra
Welwitschia
Cystopteridoideae, Diplaziopsidoideae, Rhachidosoroideae,
Seed Gymnosperms Gnetum
Conifers
Thelypteridoideae, Woodsioideae)
Euphyllophytes

Plants
ANITA grade
Pteridaceae Dennstaedtiaceae
Magnoliids
Angiosperms Monocots Lindsaeaceae Lonchitidaceae
Fabids
Cystodiaceae Saccolomataceae
Rosids Malvids

Lamiids

Asterids Campanulids

in ♀: whorls of individual megasporophylls

Cycadaceae
at trunk apex alternate with trophophylls
pachycaulous; pinnate megaphylls (no seed cones) Cycas
dioecious; males with pollen cones
motile sperm cells released within ovule
roots with N2-fixing cyanobacteria
seed cones with 2[3] seeds
per megasporophyll
Bowenia Ceratozamia Chigua Dioon
Cycadales lignins with syringaldehydes
Zamiaceae Encephalartos Lepidozamia Macrozamia
Microcycas Stangeria Zamia
spermatozoids released from dioecious; stout short shoots with

Ginkgoaceae
branched pollen tube lvs flabelliform, dichotomously veined, deciduous
acting as anchoring organ ovules 2 (basal collar); cotyledons 2 Ginkgo

Ginkgoales xeromorphic, lvs scale-like, sheathed, shed early

stems narrow, striate, photosynthetic

Ephedraceae
seed cones: 1–3 ovules; double fertilization
ovules/seeds not cotyledons 2 Ephedra
enclosed by carpel
pollen tube haustorial
primarily striate pollen caudex
anemophilous binucleate sperm lvs 2 (straplike) life-long continuous growth
dioecious venation parallel
Welwitschiaceae
primary endosperm
no bisexual strobili
"Gnetales" **
♀ gametophyte tubes grow towards pollen tubes Welwitschia
vessels
porose ectomycorrhizal
Gymnosperms

perforation lvs opposite, simple, broad, reticulate

plates

Gnetaceae
(angiosperm-like: convergence)
laticifers Gnetum
resin canals; lvs linear (needles)
monoecious
pollen mostly 2-saccate; ovules 2, inverted
Abies Cathaya Cedrus Hesperopeuce
seeds winged
Pinaceae Keteleeria Larix Nothotsuga Picea
Conifers

eustele
pollen Pinus Pseudolarix Pseudotsuga Tsuga
pollen tube lvs broad to acicular
heterospory pollen not saccate
seeds seed cones large, disintegrate
secondary growth cotyledons 2–4
Araucariaceae Agathis Araucaria Wollemia
Seed Plants P
Conifers

roots with nodules; dioecious

Acmopyle Afrocarpus Dacrycarpus Dacrydium Falcatifolium
i 1 ovule/scale, cone often one seeded
Halocarpus Lagarostrobus Lepidothamnus Manoao
Podocarpaceae
receptacle fleshy; epimatium/carpidium fleshy
n cotyledons 2
Microcachrys Nageia Parasitaxus Pherosphaera Phyllocladus
a
Spermatophytes

Podocarpus Prumnopitys Retrophyllum Saxegothaea

l monoecious
Theodor C. H. Cole, Dipl. Biol. e
Sciadopityaceae
lvs as scales; cladodes
Institute of Pharmacy and Molecular Biotechnology
s
ovules 7–9 Sciadopitys
Heidelberg University
Im Neuenheimer Feld 364
evergreen; pollen cones tiny,
D-69120 Heidelberg, Germany loss of spermatozoids
peltate microsporangiophores
Conifers

nucellar siphonogamy
1 ovule, 1 seed/cone, arillate
Amentotaxus Austrotaxus Cephalotaxus
cotyledons 2
Taxaceae Pseudotaxus Taxus Torreya
cone scales
opposite monoecious Actinostrobus Athrotaxis Austrocedrus Callitris
Prof. Dr. Hartmut H. Hilger (except Juniperus: dioecious)
Dahlem Centre of Plant Sciences (DCPS) lvs scale-like Calocedrus Chamaecyparis Cryptomeria Cunninghamia
Institute of Biology – Plant Morphology and Systematics pollen not saccate; ovules 1–20
Cupressus Diselma Glyptostrobus Fitzroya Fokienia
Cupressaceae
cotyledons many
Freie Universität Berlin
Altensteinstr. 6
seed cones terminal Juniperus Libocedrus Metasequoia Neocallitropsis
D-14195 Berlin, Germany Papuacedrus Pilgerodendron Platycladus
________________________________________________________
• hypothetical tree based on molecular phylogenetic data (July 2016)
Sequoia Sequoiadendron Taiwania Taxodium
• branch lengths deliberate, not expressing actual time scale Tetraclinis Thuja Thujopsis Widdringtonia Xanthocyparis
• 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
• References: Judd W et al. (2016); Simpson M (2010); Soltis DE et al. (2005/2011);
Christenhusz MJM et al. (2011/2014); see also: bisexual flower; fruit (ovules enclosed by carpel)
Stevens PF (2016) APweb – www.mobot.org/MOBOT/research/APweb loss of spermatozoids
* Polypodiaceae (Eupolypods I) and Aspleniaceae (Eupolypods II) each considered pollen tube penetrating stigma/style/nucellus (penetrating siphonogamy)

A ngiosperms
here sensu lato, i.e., comprising several subfamilies that in certain other

Angiosperm P hylogeny P oster

vessels; pollenkitt; primarily zoophilous
contemporary classifications are list as individual families
** the here presented sister relationship of Pinaceae to the Gnetaceae-Ephedraceae- double fertilization: triploid endosperm see
© The Authors (CC-BY)

Welwitschiaceae clade ("Gnepine"), remaining controversial to date, places the latter

within Pinales and thus Gnetales would not represent a separate order

Angiosperm Tracheophyte Bryophyte

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