Angiosperm Phylogeny

Flowering Plant Systematics

woody, vessels lacking; dioecious; flw T5–8, A∞, G5–8; 1 ovule/carpel; embryo sac 9-nucleate; 1 species (New Caledonia) A mborellales* Amborellaceae
e A g
a N r
aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A3–∞; embryo sac 4-nucleate
seeds operculate, perisperm; mucilage; alkaloids (no benzylisoquinolines) N ymphaeales * Cabombaceae Hydatellaceae Nymphaeaceae
ethereal oils in spherical idioblasts

r I a woody, vessels solitary; flw T>10, A∞, G ca.9; embryo sac 4-nucleate

l T d tiglic acid, aromatic terpenoids Austrobaileyales Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae
e lvs opposite, interpetiolar stipules; nodes swollen
y A flw small T0–3, A1–5, G1, 1 apical ovule/carpel Chloranthales* Chloranthaceae
woody; pollen uniporate
Canellales
sesquiterpenes

Canellaceae Winteraceae
a
(pellucid dots)

aromatic terpenoids

n nodes trilacunar ± herbaceous; lvs two-ranked, leaf base sheathing Aristolochiaceae Piperaceae
g
single adaxial prophyll; swollen nodes Piperales Hydnoraceae Saururaceae
i woody; lvs opposite; flw with hypanthium, staminodes frequent Calycanthaceae Hernandiaceae Monimiaceae
often valvate anthers; carpels with 1 ovule; embryo large Laurales
benzylisoquinoline alkaloids;

o Gomortegaceae Lauraceae Siparunaceae

s
Magnoliids woody; pith septate; lvs two-ranked; ovules with obturator
Annonaceae Eupomatiaceae Magnoliaceae
Magnoliales
embryo sac 8-nucleate

endosperm ruminate
p features as in
“Early Angiosperms”
Degeneriaceae Himantandraceae Myristicaceae
e infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules

r
ovules atropous, with epidermal perisperm and copious endosperm; idioblasts with ethereal oils
Acorales Acoraceae
Alismataceae (incl. Limnocharitaceae) Juncaginaceae Ruppiaceae
m mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules

s
flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green Alismatales Aponogetonaceae Butomaceae Posidoniaceae Scheuchzeriaceae
Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae
stem with ring of bundles
woody; vessels absent
eustele; nodes
sieve tube plastids
Monocots Ca oxalate
raphides
fr a follicle; East Asia Petrosaviales* Petrosaviaceae

Dioscoreales
with starch grains endosperm often twining vines; lvs often reticulate
lvs simple, persistent, entire
atactostele (scattered bundles)
no secondary thickening
nuclear
helobial, ovary often inferior, style short, branched; steroid sapogenins/alkaloids Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae
flw parts free, strobilar, mostly herbaceous nectaries
perfect, P parts varying,
often in threes,
weakly differentiated,
pollen monosulcate
sieve tube plastids with
protein crystals
when present
septal
some woody (with terminally tufted leaves)
flw spadix Pandanales Cyclanthaceae Pandanaceae Velloziaceae
stamen with broad filaments often geophytes (bulbs, tubers, rhizomes)
sympodial branching
Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae
pollen monosulcate
G apocarpous (style short in most)
nectaries absent
vascular bundles in stem scattered
lvs parallel-veined, entire
flw tepals sometimes spotted, nectaries at tepals; many seeds, seeds coat (testa) cellular
phytomelan lacking; fructans in stems, chelidonic acid, steroid saponins Liliales Colchicaceae Liliaceae Petermanniaceae Smilacaceae
no glandular teeth
embryo very small flw pentacyclic, P 3-merous, A opp. P often geophytes; anomalous sec. growth Amaryllidaceae ( incl. Agapanthaceae , Alliaceae) Iridaceae
aromatic terpenoids filaments narrow
anthers broadly attached,
septal nectary; single cotyledon
flw nonspotted tepals, septal nectaries; capsule or berry
seed coat obliterated or with phytomelan Asparagales Asparagaceae ( incl. Agavaceae, Hyacinthaceae, Ruscaceae)
Hypoxidaceae Lanariaceae Orchidaceae Tecophilaeaceae
adventitious roots
monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate Xanthorrhoeaceae ( incl. Asphodelaceae , Hemerocallidaceae)
intense primary growth, large apical meristem
infl with spathe; alkaloids Arecales Arecaceae
mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae
lvs grassy; flw often anemophilous, minute, chaffy, without nectaries Poales Cyperaceae Juncaceae Rapateaceae Typhaceae (incl. Sparganiaceae)
flw often irregular; few fertile stamens
Commelinids infl thyrsus of scorpioid cymes
phenylphenalenones Commelinales Commelinaceae Haemodoraceae Pontederiaceae
UV-fluorescing cell walls rhizomatous, large-leafed herbs; pseudostem: central infl
(ferulic/coumaric acids) flw irregular/monosymmetric, septal nectaries
Cannaceae Heliconiaceae Marantaceae Strelitziaceae
silicic acid in leaves
cuticular waxes often in rodlets
aggregated into scallops
G inferior, A often strongly modified/reduced
seeds arillate; silicic acid Zingiberales Costaceae Lowiaceae Musaceae Zingiberaceae
aquatic, herbaceous; lvs whorled, no pellucid dots; vessels lacking; monoecious
flw T9–10, A∞, G1, 1 apical ovule/carpel, pollen inaperturate, pollen tube branched; hydrophilous Ceratophyllales Ceratophyllaceae

Berberidaceae Eupteleaceae Menispermaceae

Ranunculales
lvs often divided; flw whorled, P single or multiple whorls
G apocarpous/paracarpous, superior; berberines Ranunculaceae
Circaeasteraceae Lardizabalaceae Papaveraceae

Sabiales
woody; flw 4–5-merous, nectary: disk, A epipetalous, G syncarpous
pollen colporate; endosperm helobial; triterpenoids Sabiaceae
mostly woody; flw tepals often 4-merous
A epitepalous, connectives sometimes with apical appendage Proteales Nelumbonaceae Platanaceae Proteaceae
E axial
nectary
woody; vessels lacking; idioblasts in cortex; flw tepals missing, A∞
G>5 laterally connate with abaxial nectaries; fr aggregate of follicles Trochodendrales* Trochodendraceae
u common

d mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic

Buxales* Buxaceae (incl. Didymelaceae) Haptanthaceae
benzylisoquinolines absent

flw tepals ± uniform or missing; pregnan pseudoalkaloids

i
c unisexual, dioecious; lvs toothed, sec. veins palmate
o
t
flw tepals small to lacking
ellagic acid Gunnerales Gunneraceae Myrothamnaceae
mostly woody; lvs veins proceed to apex & teeth; flw K5–∞, persisting
s
pollen tricolpate
mostly A∞, G mostly slightly connate
seeds often with aril; fr often both ventrally/dorsally dehiscent Dilleniales Dilleniaceae
flw K/C/P opp A lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent
filaments rather narrow Altingiaceae Daphniphyllaceae Hamamelidaceae
Saxifragales
fr mostly dry, dehiscent
anthers basifixed
nodes trilacunar
myricetin, flavonols Cercidiphyllaceae Grossulariaceae Paeoniaceae
stomata anomocytic
often tendrillar vines; lvs often divided and with glandular teeth Crassulaceae Haloragaceae Saxifragaceae
ethereal oils absent

Vitales*
A epipetalous, 2 ovules per carpel; raphides, pearl glands
berries Vitaceae
cork origin deep-seated

Zygophyllales*
endosperm lacking
resinous, lignans/neolignans, harman alkaloids Krameriaceae Zygophyllaceae
ellagic acid tannins

flw small, G often 3-merous, nectary: intrastaminal disk

seeds often with aril (red-orange)
infl cymose
lvs often compound, pulvini (sleep movement)
flw A5 or multiple, branched style common
mucilage cells; oxalates
Celastraceae Lepidobotryaceae
Celastrales (incl. Hippocrateaceae, Brexiaceae) Parnassiaceae
Brunelliaceae Connaraceae Elaeocarpaceae
Oxalidales
Oxalidaceae
Cephalotaceae Cunoniaceae Huaceae

F Achariaceae Euphorbiaceae Ochnaceae Podostemaceae

a lvs margins toothed
flw G often tricarpellate Malpighiales Chrysobalanaceae
Clusiaceae
Hypericaceae
Linaceae
Passifloraceae
Phyllanthaceae
Rhizophoraceae
Salicaceae
b
i flw often “papilionaceous”: wing, standard, keel, mostly G1 Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae
mostly A10; fr a pod; symbiosis with root nodule bacteria
d diverse alkaloids, NP amino acids, lectins (in Fabaceae) Fabales Fabaceae Polygalaceae Quillajaceae Surianaceae
c E s
Barbeyaceae Elaeagnaceae Rosaceae
o u lvs mostly simple with stipules

Rosales
flw K valvate (and hypanthium) persisting
r d N fix
carpels with 1 ovule, stigma dry; dihydroflavonols Cannabaceae Moraceae Ulmaceae
e i embryo large, Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae)
c lvs with
stipules
endosperm scanty lvs mostly alternate
Anisophyllaceae Coriariaceae Cucurbitaceae
Cucurbitales
flw often unisexual, G mostly inferior
o parietal placentation; cucurbitacins Tetramelaceae
t Begoniaceae Corynocarpaceae Datiscaceae
s R lvs undivided; flw small, unisexual
o anemophilous, thus T reduced or lacking, G mostly inferior
Betulaceae Fagaceae Myricaceae Rhoipteleaceae
flw pentacyclic s
infl spikes or catkins; fr 1-seeded, mostly nuts
ectomycorrhiza; tannins, dihydroflavonols Fagales Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae
parts alternating i lvs with glandular teeth

Geraniales
G connate
K + C (free) d flw A obdiplostemonous, nectary on filament
Francoaceae Geraniaceae Ledocarpaceae Melianthaceae
stems jointed at nodes; ethereal oils, ellagic acid
(A polyandrous) s
pollen tricolporate
trihydroxyl-flavonoids
lvs opposite, colleters (glandular hair on adaxial surface of petiole base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae)
stipules small (if any); cork deep seated
Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae)
flw K valvate, persisting, A incurved in bud, ovary inferior; ovules many
endosperm scanty; scaly bark; flavonols, myricetin Myrtales Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae
woody
Crossosomataceae Stachyuraceae
K + C, stylodia free
hypanthium, nectary disk Crossosomatales Geissolomataceae Staphyleaceae
Strasburgeriaceae
trees; lvs spiral; extrafloral nectaries
M staminate flw: A = and opposite C
bark bitter, anthraquinones Picramniales* Picramniaceae
a mostly woody (silica/silicified)
l Anacardiaceae Meliaceae Rutaceae
v
lvs alternate, odd-pinnately compound
flw often imperfect, intrastaminal disk; ethereal oils Sapindales Burseraceae Nitrariaceae Sapindaceae
Simaroubaceae

i vessel elements: scalariform perforations; mucilage cells
d embryo
lvs margins toothed, stipules cauline
flw small, A = and opposite K, ovules 1-2/carpel Huerteales* Dipentodontaceae Gerrardinaceae Petenaeaceae Tapisciaceae
s large
bark fibrous; hairs often stellate Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
endo-
sperm flw K often valvate
scanty petals contorted, A often ∞; mucilage
cyclopropenoid fatty acids, flavones
Malvales Cistaceae
Dipterocarpaceae
Cytinaceae
Neuradaceae
Muntingiaceae
Sphaerosepalaceae
Sarcolaenaceae
Thymelaeaceae
lvs alternate; flw often 4-merous
often clawed petals, A often many, 2×K Bataceae Caricaceae Limnanthaceae Salvadoraceae
hypogynous (often gynophore); infl racemose
myrosin cells, glucosinolates Brassicales Brassicaceae Cleomaceae Moringaceae Tovariaceae
woody; parasites or semiparasites; without mycorrhiza Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae
Liverworts
lvs margin entire; flw A epipetalous
Balanophoraceae Misodendraceae Opiliaceae Schoepfiaceae
Mosses perianth often simple, valvate, persisting
polyacetylenes, triterpene sapogenins, silicic acid Santalales Loranthaceae Olacaceae Santalaceae (incl. Viscaceae)
Hornworts woody; lvs stomata cyclocytic,

Berberidopsidales* Aextoxicaceae Berberidopsidaceae

petiole bundles annular; fr fleshy
Lycophytes calcium oxalate as crystals
Ferns mostly herbaceous; without mycorrhiza Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae
(incl. horsetails)
G often unilocular with central placentation, Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae
Palmferns
Ginkgo
pollen colpate, surface spiny
betalains or anthocyanins (latter, e.g., in Caryophyllaceae) Caryophyllales [incl. Chenopodiaceae] Droseraceae
Basellaceae Drosophyllaceae
Nyctaginaceae
Phytolaccaceae
Simmondsiaceae
Talinaceae
Ephedra
Welwitschia mostly woody; lvs mostly undivided, hydathode teeth Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae
Seed Gymnosperms Gnetum flw often 4-merous, K much smaller than C, persisting,
Plants intrastaminal disk, G inferior; fr drupaceous
Cornales Cornaceae Grubbiaceae Loasaceae
Conifers
diverse iridoids
ANITA grade Curtisiaceae Hydrangeaceae Nyssaceae
Magnoliids
lvs teeth, theoid; nodes unilacunar Actinidiaceae Ericaceae Polemoniaceae Sarraceniaceae
Angiosperms flw 5-merous, pentacyclic
Monocots Balsaminaceae Fouquieriaceae Primulaceae Styracaceae
Fabids
nonhydrolyzable tannins, ellagic acid, hydroquinones
Ericales Clethraceae Lecythidaceae Roridulaceae Theaceae
Malvids
woody; dioecious Ebenaceae Myrsinaceae§ Sapotaceae Theophrastaceae
Rosids flw small, C valvate, G unilocular; fruit indehiscent
Lamiids
A
iridoids (aucubin), gutta
Garryales Eucommiaceae Garryaceae (incl. Aucubaceae)
lvs opposite, colleters
Campanulids s Apocynaceae (incl. Asclepiadaceae) Loganiaceae
Gentianales
Asterids flw corolla convolute in bud
t late
sympetaly
indole alkaloids; iridoids Gentianaceae Gelsemiaceae Rubiaceae
e lvs opposite; nodes 1:1; flw mostly monosymmetric Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae
r
L
A often 2(+2); ethereal oils in gland-headed hairs
Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae
i
d a
6-oxygenated flavones, rosmarinic acid,
oligosaccharides: cornoside, verbascoside (acetoside) Lamiales Byblidaceae Martyniaceae Pedaliaceae Stilbaceae
Theodor C. H. Cole, Dipl. Biol.
s A=C,
m lvs spiral, simple; nodes unilacunar
Gesneriaceae Hydrostachyaceae Oleaceae Phrymaceae Verbenaceae
Institut für Pharmazie und ii
Molekulare Biotechnologie mostly sympetalous
epipetalous
polyandry
rare, d
flw petals plicate; K persisting
diverse alkaloids, no iridoids Solanales Convolvulaceae (incl. Cuscutaceae)
Hydroleaceae Montiniaceae
Solanaceae (incl. Nolanaceae)
Sphenocleaceae
Universität Heidelberg nectary gynoecial

Im Neuenheimer Feld 364

ovules unitegmic G(2) s lvs roughly hairy; nodes unilacunar
Boraginaceae Cordiaceae Heliotropiaceae
Boraginales
endosperm cellular infl infl scorpioid; mostly 4 ovules
D-69120 Heidelberg, Germany iridoids common cymose isokestose, higher inulins, pyrrolizidine alkaloids Wellstediaceae
ellagic acid Codonaceae Ehretiaceae (+ Lennoa) Hydrophyllaceae
lacking woody; lvs serrate;

early
flw 1-2 ovules/carpel, C± free, K slightly connate
fr drupe with broad stigma Aquifoliales Aquifoliaceae Cardiopteridaceae Stemonuraceae
Prof. Dr. Hartmut H. Hilger sympetaly
flw often monosymmetric
Institut für Biologie
flw small
plunger pollination (long style)
Asteraceae Goodeniaceae Pentaphragmataceae
Asterales
embryo
Botanik – Morphologie und Systematik short nodes trilacunar Calyceraceae Menyanthaceae Rousseaceae
inulin, sesquiterpenes, secoiridoids
Freie Universität Berlin Campanulaceae (incl. Lobeliaceae) Stylidiaceae
Altensteinstr. 6
D-14195 Berlin, Germany C mostly woody; infl racemose, C free
anthers basifixed, nectary disc Escalloniales* Escalloniaceae
a
m woody, evergreen; nodes 1:1
flw polysymmetric, anthers basifixed Bruniales* Bruniaceae Columelliaceae (incl. Desfontainia)
p
G inferior

_______________________________________________________________________________________ woody (except Apiaceae)

- hypothetical tree based on molecular phylogenetic data (Jan 2012); branch lengths deliberate, not expressing actual time scale a lvs often divided; nodes usu. multilacunar Apiaceae Griseliniaceae Pennantiaceae
- 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)
n
u
infl mostly umbel; drupe or schizocarp
(Apiaceae: mericarp/carpophore)
A piales Araliaceae Myodocarpaceae Pittosporaceae
- this poster depicts only the largest and most important of the currently accepted approx. 450 families (according to APweb 2012)
1–few-seeded

l
fr usually

woody; infl racemose, flw 4-merous,

Paracryphiales*
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); i filaments stout, capsule septicidal Paracryphiaceae
Stevens PF (2012) APweb – www.mobot.org/MOBOT/research/APweb; Watson/Dallwitz (2009) delta-intkey.com/angio/
d lvs opposite, often basally connate
§ Myrsinaceae and Theophrastaceae again included in Primulaceae by APG III
s nodes 3:3; buds with scales
Adoxaceae Diervillaceae Linnaeaceae
Thanks to Christoph Dobeš, Ray F. Evert, Peter H. Raven, Douglas E. Soltis, Peter F. Stevens, Maximilian Weigend, Michael Wink
flw often monosymmetric
K persistent in fruit; secoiridoids Dipsacales Caprifoliaceae Dipsacaceae Morinaceae
Valerianaceae