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
Lycophytes

lvs ligulate sporangia in axils of sporophylls

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
Aspleniaceae
Mosses (incl. Asplenoideae, Athyrioideae, Blechnoideae,
sporangial maturation mixed
Hornworts Cystopteridoideae, Diplaziopsidoideae, Rhachidosoroideae,
Polypodiales
sporangium with vertical annulus
euphyll Lycophytes
interrupted by stalk and stomium Thelypteridoideae, Woodsioideae)
leaf gaps
protoxylem exarch Ferns
roots monopodial
(incl. horsetails)
Polypodiaceae
30-kb cp inversion Palmferns
Ginkgo (incl. Polypodioideae, Davallioideae,
Ephedra Didymochlaenoideae, Dryopteridoideae, Hypodematioideae,
Welwitschia
Seed Gymnosperms Gnetum Lomariopsidoideae, Oleandroideae, Tectarioideae)
Conifers
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
vessels
♀ gametophyte tubes grow towards pollen tubes Welwitschia
porose
perforation
ectomycorrhizal
plates
Gymnosperms

lvs opposite, simple, broad, reticulate

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 Pinus Picea
Conifers

eustele
pollen 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
Spermatophytes

Theodor C. H. Cole, Dipl. Biol. a

Podocarpus Prumnopitys Retrophyllum Saxegothaea
Institute of Pharmacy and Molecular Biotechnology
Heidelberg University
l monoecious
e
Sciadopityaceae
Im Neuenheimer Feld 364 lvs as scales; cladodes
D-69120 Heidelberg, Germany s
ovules 7–9 Sciadopitys
evergreen; pollen cones tiny,
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
Prof. Dr. Hartmut H. Hilger opposite Actinostrobus Athrotaxis Austrocedrus Callitris
monoecious (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
Freie Universität Berlin
Altensteinstr. 6
cotyledons many
seed cones terminal Cupressaceae Juniperus Libocedrus Metasequoia Neocallitropsis
D-14195 Berlin, Germany Papuacedrus Pilgerodendron Platycladus
Sequoia Sequoiadendron Taiwania Taxodium
Tetraclinis Thuja Thujopsis Widdringtonia Xanthocyparis
_______________________________________________________
• hypothetical tree based on molecular phylogenetic data (Feb. 2014)
• branch lengths deliberate, not expressing actual time scale position bisexual flower; fruit (ovules enclosed by carpel)
of many characters on tree unclear loss of spermatozoids
• if a character is marked as being a potential synapomorphy at a node/for a clade, pollen tube penetrating stigma/style/nucellus (penetrating siphonogamy)

A ngiosperms
this does not mean that all members of that clade possess that character

Angiosperm P hylogeny P oster

vessels; pollenkitt; primarily zoophilous;
• References: Judd W et al. (2007); Simpson M (2010); Soltis DE et al. (2005/2011);
Christenhusz MJM et al. (2011/2014)
double fertilization: triploid endosperm see
see also: Stevens PF (2011) APweb – www.mobot.org/MOBOT/research/APweb

Angiosperm Tracheophte Bryophte

Phylogeny Phylogeny Phylogeny
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