Angiosperm Phylogeny
Flowering Plant Systematics
woody, vessels lacking; dioecious; flw T58, A, G58; 1 ovule/carpel; embryo sac 9-nucleate; 1 species (New Caledonia)
aquatic, herbaceous; cambium absent; aerenchyma; flw T412, A3; embryo sac 4-nucleate
seeds operculate, perisperm; mucilage; alkaloids (no benzylisoquinolines)
r
a
woody, vessels solitary; flw T>10, A, G ca.9; embryo sac 4-nucleate
tiglic acid, aromatic terpenoids
d
e
lvs opposite, interpetiolar stipules; nodes swollen
flw small T03, A15, G1, 1 apical ovule/carpel
woody; pollen uniporate
aromatic terpenoids
sesquiterpenes
ethereal oils in spherical idioblasts
(pellucid dots)
e
a
r
l
y
A
N
I
T
A
benzylisoquinoline alkaloids;
embryo sac 8-nucleate
n
g
i
o
s
p
e
r
m
s
nodes trilacunar
woody; lvs opposite; flw with hypanthium, staminodes frequent
often valvate anthers; carpels with 1 ovule; embryo large
Magnoliids
woody; pith septate; lvs two-ranked; ovules with obturator
endosperm ruminate
features as in
Early Angiosperms
infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules
ovules atropous, with epidermal perisperm and copious endosperm; idioblasts with ethereal oils
mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules
flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green
Monocots
stem with ring of bundles
fr a follicle; East Asia
Ca oxalate
raphides
endosperm
nuclear
helobial,
nectaries
atactostele (scattered bundles)
no secondary thickening
mostly herbaceous
pollen monosulcate
sieve tube plastids with
protein crystals
sympodial branching
vascular bundles in stem scattered
lvs parallel-veined, entire
no glandular teeth
flw pentacyclic, P 3-merous, A opp. P
filaments narrow
anthers broadly attached,
septal nectary; single cotyledon
adventitious roots
woody; vessels absent
eustele; nodes
sieve tube plastids
with starch grains
lvs simple, persistent, entire
flw parts free, strobilar,
perfect, P parts varying,
often in threes,
weakly differentiated,
stamen with broad filaments
pollen monosulcate
G apocarpous (style short in most)
nectaries absent
embryo very small
aromatic terpenoids
herbaceous; lvs two-ranked, leaf base sheathing
single adaxial prophyll; swollen nodes
often twining vines; lvs often reticulate
ovary often inferior, style short, branched; steroid sapogenins/alkaloids
some woody (with terminally tufted leaves)
flw spadix
when present
septal
often geophytes (bulbs, tubers, rhizomes)
flw tepals sometimes spotted, nectaries at tepals; many seeds, seeds coat (testa) cellular
phytomelan lacking; fructans in stems, chelidonic acid, steroid saponins
often geophytes; anomalous sec. growth
capsule or berry
seed coat obliterated or with phytomelan
monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate
intense primary growth, large apical meristem
infl with spathe; alkaloids
mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent
lvs grassy; flw often anemophilous, minute, chaffy, without nectaries
flw often irregular; few fertile stamens
infl thyrsus of scorpioid cymes
phenylphenalenones
Commelinids
UV-fluorescing cell walls
(ferulic/coumaric acids)
silicic acid in leaves
rhizomatous, large-leafed herbs; pseudostem: central infl
flw irregular/monosymmetric, septal nectaries
G inferior, A often strongly modified/reduced
seeds arillate; silicic acid
cuticular waxes often in rodlets
aggregated into scallops
aquatic, herbaceous; lvs whorled, no pellucid dots; vessels lacking; monoecious
flw T910, A, G1, 1 apical ovule/carpel, pollen inaperturate, pollen tube branched; hydrophilous
lvs often divided; flw whorled, P single or multiple whorls
G apocarpous/paracarpous, superior; berberines
woody; flw 45-merous, nectary: disk, A epipetalous, G syncarpous
pollen colporate; endosperm helobial; triterpenoids
mostly woody; flw tepals often 4-merous
A epitepalous, connectives sometimes with apical appendage
u
d
i
c
o
t
s
woody; vessels lacking; idioblasts in cortex; flw tepals missing, A
G>5 laterally connate with abaxial nectaries; fr aggregate of follicles
axial
nectary
common
mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic
flw tepals uniform or missing; pregnan pseudoalkaloids
benzylisoquinolines absent
pollen tricolpate
flw K/C/P opp A
filaments rather narrow
anthers basifixed
nodes trilacunar
stomata anomocytic
ethereal oils absent
unisexual, dioecious; lvs toothed, sec. veins palmate
flw tepals small to lacking
ellagic acid
mostly woody; lvs veins proceed to apex & teeth; flw K5, persisting
mostly A, G mostly slightly connate
seeds often with aril; fr often both ventrally/dorsally dehiscent
lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent
fr mostly dry, dehiscent
myricetin, flavonols
often tendrillar vines; lvs often divided and with glandular teeth
A epipetalous, 2 ovules per carpel; raphides, pearl glands
berries
ellagic acid tannins
stipules;
nodes
3:3
c
o
r
e
cork origin deep-seated
endosperm lacking
resinous, lignans/neolignans, harman alkaloids
flw small, G often 3-merous, nectary: intrastaminal disk
seeds often with aril (red-orange)
infl cymose
o
s
i
d
s
lvs often compound, pulvini (sleep movement)
flw A5 or multiple, branched style common
mucilage cells; oxalates
lvs margins toothed
flw G often tricarpellate
a
b
i
d
s
u
d
i
c
o
t
s
flw often papilionaceous: wing, standard, keel, mostly G1
mostly A10; fr a pod; symbiosis with root nodule bacteria
diverse alkaloids, NP amino acids, lectins (in Fabaceae)
lvs mostly simple with stipules
flw K valvate (and hypanthium) persisting
carpels with 1 ovule, stigma dry; dihydroflavonols
N fix
embryo large,
endosperm scanty
lvs mostly alternate
flw often unisexual, G mostly inferior
parietal placentation; cucurbitacins
lvs undivided; flw small, unisexual
anemophilous, thus T reduced or lacking, G mostly inferior
infl spikes or catkins; fr 1-seeded, mostly nuts
ectomycorrhiza; tannins, dihydroflavonols
flw pentacyclic
parts alternating
G connate
K + C (free)
(A polyandrous)
pollen tricolporate
trihydroxyl-flavonoids
lvs with glandular teeth
flw A obdiplostemonous, nectary on filament
stems jointed at nodes; ethereal oils, ellagic acid
lvs opposite, colleters (glandular hair on adaxial surface of petiole base)
stipules small (if any); cork deep seated
flw K valvate, persisting, A incurved in bud, ovary inferior; ovules many
endosperm scanty; scaly bark; flavonols, myricetin
woody
K + C, stylodia free
hypanthium, nectary disk
trees; lvs spiral; extrafloral nectaries
staminate flw: A = and opposite C
bark bitter, anthraquinones
M
a
l
v
i
d
s
mostly woody (silica/silicified)
lvs alternate, odd-pinnately compound
flw often imperfect, intrastaminal disk; ethereal oils
vessel elements: scalariform perforations; mucilage cells
lvs margins toothed, stipules cauline
flw small, A = and opposite K, ovules 1-2/carpel
embryo
large
bark fibrous; hairs often stellate
flw K often valvate
petals contorted, A often ; mucilage
cyclopropenoid fatty acids, flavones
endosperm
scanty
Liverworts
lvs alternate; flw often 4-merous
often clawed petals, A often many, 2K
hypogynous (often gynophore); infl racemose
myrosin cells, glucosinolates
woody; parasites / semiparasites; without mycorrhiza
lvs margin entire; flw A epipetalous
perianth often simple, valvate, persisting
polyacetylenes, triterpene sapogenins, silicic acid
Mosses
Hornworts
Lycophytes
Ferns
(incl. horsetails)
woody; lvs stomata cyclocytic,
petiole bundles annular; fr fleshy
calcium oxalate as crystals
Palmferns
Ginkgo
Ephedra
Welwitschia
Gnetum
Conifers
Seed Gymnosperms
Plants
mostly herbaceous; without mycorrhiza
G often unilocular with central placentation,
pollen colpate, surface spiny
betalains or anthocyanins (latter, e.g., in Caryophyllaceae)
ANITA grade
Magnoliids
Angiosperms
mostly woody; lvs mostly undivided, hydathode teeth
flw often 4-merous, K much smaller than C, persisting,
intrastaminal disk, G inferior; fr drupaceous
diverse iridoids
Monocots
Fabids
Rosids
Malvids
lvs teeth, theoid; nodes unilacunar
flw 5-merous, pentacyclic
nonhydrolyzable tannins, ellagic acid, hydroquinones
Lamiids
Campanulids
A
s
t
e
r
i
d
s
Theodor C. H. Cole, Dipl. Biol.
Institute of Pharmacy and Molecular Biotechnology
Heidelberg University
Im Neuenheimer Feld 364
D-69120 Heidelberg, Germany
mostly sympetalous
nectary gynoecial
ovules unitegmic
endosperm cellular
iridoids common
Prof. Dr. Hartmut H. Hilger
Dahlem Centre of Plant Sciences (DCPS)
Institute of Biology Plant Morphology and Systematics
Freie Universitt Berlin
Altensteinstr. 6, D-14195 Berlin, Germany
late
sympetaly
lvs spiral, simple; nodes unilacunar
flw petals plicate; K persisting
diverse alkaloids, no iridoids
A=C,
epipetalous
polyandry
rare,
G(2)
a
m
ii
d
s
lvs opposite; nodes 1:1; flw mostly monosymmetric
A often 2(+2); ethereal oils in gland-headed hairs
6-oxygenated flavones, rosmarinic acid,
oligosaccharides: cornoside, verbascoside (acetoside)
lvs roughly hairy; nodes unilacunar
infl scorpioid; mostly 4 ovules
isokestose, higher inulins, pyrrolizidine alkaloids
infl
cymose
ellagic acid
lacking
woody; lvs serrate;
flw 1-2 ovules/carpel, C free, K slightly connate
fr drupe with broad stigma
Myrsinaceae and Theophrastaceae again included in Primulaceae by APG III
Thanks to Christoph Dobe, Ray F. Evert, Marc Gottschling, Richard G. Olmstead, Peter H. Raven, Douglas E. Soltis, Peter F. Stevens, Maximilian Weigend, Michael Wink
Bryophte
Phylogeny
Poster
flw often monosymmetric
plunger pollination (long style)
nodes trilacunar
inulin, sesquiterpenes, secoiridoids
G inferior
a
m
p
a
n
u
l
i
d
s
mostly woody; infl racemose, C free
anthers basifixed, nectary disc
fr usually
- hypothetical tree based on molecular phylogenetic data (Feb. 2014); branch lengths deliberate, not expressing actual time scale
- position of many characters on tree unclear; 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; * orders added as of APG III (2009)
- this poster depicts only the largest and most important of the currently accepted approx. 450 families (according to APweb 2014);
for family characteristics see: Kubitzki K, ed. (1990 ff).
- References: APG III (2009); Judd W et al. (2007); Simpson M (2010); Soltis DE et al. (2005/2011/2014);
Stevens PF (2014) APweb www.mobot.org/MOBOT/research/APweb; Watson/Dallwitz (2009) delta-intkey.com/angio/
Tracheophte
Phylogeny
Poster
lvs opposite, colleters
flw corolla convolute in bud
indole alkaloids; iridoids
early
sympetaly
flw small
embryo
short
________________________________________________________________________________________
Angiosperm
Phylogeny
Poster
woody; dioecious
flw small, C valvate, G unilocular; fruit indehiscent
iridoids (aucubin), gutta
1few-seeded
Asterids
woody, evergreen; nodes 1:1
flw polysymmetric, anthers basifixed
woody (except Apiaceae)
lvs often divided; nodes usu. multilacunar
infl mostly umbel; drupe or schizocarp
(Apiaceae: mericarp/carpophore)
woody; infl racemose, flw 4-merous,
filaments stout, capsule septicidal
lvs opposite, often basally connate
nodes 3:3; buds with scales
flw often monosymmetric
K persistent in fruit; secoiridoids
A mborellales*
N ymphaeales *
Amborellaceae
Cabombaceae
Hydatellaceae
Austrobaileyales
Austrobaileyaceae
Chloranthales*
Chloranthaceae
Canellales
Canellaceae
Winteraceae
Aristolochiaceae
Hydnoraceae
Piperaceae
Saururaceae
Piperales
Laurales
Calycanthaceae
Gomortegaceae
Magnoliales
Annonaceae
Degeneriaceae
Acorales
Acoraceae
Nymphaeaceae
Schisandraceae (incl. Illiciaceae)
Hernandiaceae
Lauraceae
Monimiaceae
Siparunaceae
Eupomatiaceae
Himantandraceae
Magnoliaceae
Myristicaceae
Alismatales
Alismataceae (incl. Limnocharitaceae) Juncaginaceae
Aponogetonaceae
Butomaceae
Posidoniaceae
Araceae
Hydrocharitaceae Potamogetonaceae
Petrosaviales*
Petrosaviaceae
Dioscoreales
Pandanales
Burmanniaceae Dioscoreaceae
Nartheciaceae
Cyclanthaceae
Velloziaceae
Liliales
Alstroemeriaceae
Colchicaceae
Asparagales
Arecales
Trimeniaceae
Pandanaceae
Corsiaceae
Liliaceae
Ruppiaceae
Scheuchzeriaceae
Zosteraceae
Taccaceae
Melanthiaceae
Petermanniaceae
Philesiaceae
Smilacaceae
Amaryllidaceae ( incl. Agapanthaceae , Alliaceae) Iridaceae
Asparagaceae ( incl. Agavaceae, Hyacinthaceae, Ruscaceae)
Hypoxidaceae Lanariaceae Orchidaceae Tecophilaeaceae
Xanthorrhoeaceae ( incl. Asphodelaceae , Hemerocallidaceae)
Arecaceae
Poales
Bromeliaceae Eriocaulaceae Poaceae Restionaceae
Xyridaceae
Cyperaceae Juncaceae Rapateaceae Typhaceae (incl. Sparganiaceae)
Commelinales
Commelinaceae
Zingiberales
Cannaceae
Costaceae
Ceratophyllales
Ceratophyllaceae
Ranunculales
Sabiales
Proteales
Berberidaceae
Circaeasteraceae
Trochodendrales*
Trochodendraceae
Buxales*
Buxaceae Didymelaceae
Gunnerales
Gunneraceae Myrothamnaceae
Dilleniales
Dilleniaceae
Saxifragales
Altingiaceae Daphniphyllaceae
Cercidiphyllaceae
Grossulariaceae
Crassulaceae
Haloragaceae
Vitales*
Haemodoraceae
Heliconiaceae
Lowiaceae
Pontederiaceae
Marantaceae
Musaceae
Eupteleaceae
Lardizabalaceae
Strelitziaceae
Zingiberaceae
Menispermaceae
Papaveraceae
Ranunculaceae
Sabiaceae
Nelumbonaceae
Platanaceae
Proteaceae
Hamamelidaceae
Paeoniaceae
Saxifragaceae
Vitaceae
Zygophyllales*
Krameriaceae
Celastrales
Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae)
Lepidobotryaceae
Oxalidales
Brunelliaceae
Cephalotaceae
Malpighiales
Achariaceae
Euphorbiaceae
Rafflesiaceae
Ochnaceae
Chrysobalanaceae
Hypericaceae
Passifloraceae
Clusiaceae
Linaceae
Phyllanthaceae
Erythroxylaceae
Malpighiaceae Picrodendraceae
Fabales
Zygophyllaceae
Connaraceae
Cunoniaceae
Elaeocarpaceae
Oxalidaceae
Huaceae
Podostemaceae
Rhizophoraceae
Salicaceae
Violaceae
Fabaceae Polygalaceae Quillajaceae Surianaceae
Rosales
Barbeyaceae
Cannabaceae
Dirachmaceae
Elaeagnaceae
Moraceae
Rhamnaceae
Cucurbitales
Anisophyllaceae
Begoniaceae
Coriariaceae
Corynocarpaceae
Fagales
Betulaceae
Fagaceae
Casuarinaceae Juglandaceae
Geraniales
Francoaceae
Rosaceae
Ulmaceae
Urticaceae (incl. Cecropiaceae)
Cucurbitaceae
Datiscaceae
Myricaceae
Nothofagaceae
Geraniaceae
Tetramelaceae
Rhoipteleaceae
Ticodendraceae
Ledocarpaceae
Melianthaceae
Myrtales
Combretaceae
Myrtaceae
Penaeaceae (incl. Oliniaceae)
Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae)
Melastomataceae (incl. Memecylaceae)
Onagraceae
Vochysiaceae
Crossosomatales
Crossosomataceae Stachyuraceae
Strasburgeriaceae
Geissolomataceae
Staphyleaceae
Picramniales*
Picramniaceae
Sapindales
Anacardiaceae
Burseraceae
Huerteales*
Dipentodontaceae
Meliaceae
Nitrariaceae
Gerrardinaceae
Malvales
Bixaceae
Cistaceae
Dipterocarpaceae
Brassicales
Bataceae
Brassicaceae
Capparaceae
Santalales
Balanophoraceae
Loranthaceae
Misodendraceae
Olacaceae
Berberidopsidales*
Aextoxicaceae
Berberidopsidaceae
Caryophyllales
Simaroubaceae
Petenaeaceae
Tapisciaceae
Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
Cytinaceae
Muntingiaceae
Sarcolaenaceae
Neuradaceae
Sphaerosepalaceae
Thymelaeaceae
Caricaceae
Cleomaceae
Koeberliniaceae
Limnanthaceae
Moringaceae
Resedaceae
Aizoaceae
Amaranthaceae
Caryophyllaceae
Didiereaceae
[incl. Chenopodiaceae] Droseraceae
Basellaceae
Drosophyllaceae
Cactaceae
Frankeniaceae
Cornales
Cornaceae
Curtisiaceae
Ericales
Actinidiaceae
Balsaminaceae
Clethraceae
Ebenaceae
Ericaceae
Fouquieriaceae
Lecythidaceae
Myrsinaceae
Eucommiaceae
Garryales
Rutaceae
Sapindaceae
Opiliaceae
Schoepfiaceae
Santalaceae (incl. Viscaceae)
Molluginaceae
Nepenthaceae
Nyctaginaceae
Phytolaccaceae
Plumbaginaceae
Grubbiaceae
Hydrangeaceae
Salvadoraceae
Tovariaceae
Tropaeolaceae
Polygonaceae
Portulacaceae
Simmondsiaceae
Talinaceae
Tamaricaceae
Loasaceae
Nyssaceae
Polemoniaceae
Sarraceniaceae
Primulaceae
Styracaceae
Roridulaceae
Theaceae
Sapotaceae
Theophrastaceae
Garryaceae (incl. Aucubaceae)
Gentianales
Apocynaceae (incl. Asclepiadaceae)
Loganiaceae
Gentianaceae Gelsemiaceae Rubiaceae
Solanales
Convolvulaceae (incl. Cuscutaceae)
Solanaceae (incl. Nolanaceae)
Hydroleaceae
Montiniaceae
Sphenocleaceae
Acanthaceae
Lamiaceae
Orobanchaceae
Plantaginaceae
Bignoniaceae
Lentibulariaceae Paulowniaceae
Scrophulariaceae
Byblidaceae
Martyniaceae
Pedaliaceae
Stilbaceae
Gesneriaceae Hydrostachyaceae Oleaceae Phrymaceae Verbenaceae
Lamiales
Boraginales
Boraginaceae
Codonaceae
Ehretiaceae (+ Lennoaceae) Cordiaceae
Aquifoliales
Aquifoliaceae
Asterales
Asteraceae Goodeniaceae Pentaphragmataceae
Calyceraceae
Menyanthaceae
Rousseaceae
Campanulaceae (incl. Lobeliaceae)
Stylidiaceae
Escalloniales*
Wellstediaceae
Heliotropiaceae
Cardiopteridaceae
Hydrophyllaceae
(Namaceae)
Stemonuraceae
Escalloniaceae
Bruniales*
Bruniaceae
A piales
Apiaceae
Griseliniaceae
Pennantiaceae
Araliaceae Myodocarpaceae Pittosporaceae
Paracryphiales*
Paracryphiaceae
Dipsacales
Adoxaceae
Caprifoliaceae
Columelliaceae (incl. Desfontainia)
Diervillaceae
Dipsacaceae
Linnaeaceae
Morinaceae
Valerianaceae
