Angiosperm Phylogeny

Flowering Plant Systematics

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

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

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

sesquiterpenes
Canellales Canellaceae Winteraceae
(pellucid dots)
Early Angiosperms

aromatic terpenoids

nodes trilacunar ± herbaceous; lvs two-ranked, leaf base sheathing; Aristolochiaceae Piperaceae
single adaxial prophyll; swollen nodes Piperales Hydnoraceae Saururaceae
benzylisoquinoline alkaloids;

woody; lvs opposite; flw with hypanthium, staminodes frequent, Calycanthaceae Hernandiaceae Monimiaceae
often valvate anthers; carpels with 1 ovule; embryo large
Laurales Gomortegaceae Lauraceae Siparunaceae
Magnoliids
embryo sac 8-nucleate

woody; pith septate; lvs two-ranked; ovules with obturator;

Annonaceae Eupomatiaceae Magnoliaceae
features as in
“Early Angiosperms”
endosperm ruminate Magnoliales Degeneriaceae Himantandraceae Myristicaceae
infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules;
ovules atropous, with epidermal perisperm and copious endosperm; idioblasts with ethereal oils
Acorales Acoraceae
mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules; Alismataceae (incl. Limnocharitaceae) Juncaginaceae Ruppiaceae
flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green Alismatales Aponogetonaceae Butomaceae Posidoniaceae Scheuchzeriaceae
woody; vessels absent, Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae
eustele; nodes; stem with ring of bundles;
sieve tube plastids
with starch grains; Monocots Ca oxalate
raphides
fr a follicle; East Asia Petrosaviales* Petrosaviaceae
lvs simple, persistent, entire;
Dioscoreales
endosperm often twining vines; lvs often reticulate;
flw parts free, strobilar, atactostele (scattered bundles), nuclear
ovary often inferior, style short, branched; steroid sapogenins/alkaloids Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae
perfect, P parts varying, no secondary thickening; helobial,
often in threes, mostly herbaceous; nectaries
weakly differentiated,
stamen with broad filaments,
pollen monosulcate;
sieve tube plastids with
when present
septal
some woody (with terminally tufted leaves),
flw spadix Pandanales Cyclanthaceae Pandanaceae Velloziaceae
pollen monosulcate; protein crystals;
often geophytes (bulbs, tubers, rhizomes);
G apocarpous sympodial branching; Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae
(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
embryo very small; no glandular teeth;
aromatic terpenoids flw pentacyclic, P 3-merous, A opp. P, often geophytes; anomalous sec. growth; Amaryllidaceae ( incl. Agapanthaceae , Alliaceae)
Asparagales
filaments narrow,
anthers broadly attached,
flw nonspotted tepals, septal nectaries; capsule or berry; Asparagaceae ( incl. Agavaceae, Hyacinthaceae, Ruscaceae)
seed coat obliterated or with phytomelan
septal nectary; single cotyledon; Hypoxidaceae Iridaceae Lanariaceae Orchidaceae
adventitious roots
monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate; Xanthorrhoeaceae ( incl. Asphodelaceae , Hemerocallidaceae)
Arecales
intense primary growth, large apical meristem;
infl with spathe; alkaloids 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,
silicic acid in leaves; Cannaceae Heliconiaceae Marantaceae Strelitziaceae
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
lvs often divided; flw whorled, P single or multiple whorls,
Berberidaceae Eupteleaceae Menispermaceae
G apocarpous/paracarpous, superior;
berberines Ranunculales Circaeasteraceae Lardizabalaceae Papaveraceae
Ranunculaceae

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
woody; vessels lacking; idioblasts in cortex; flw tepals missing, A∞,
Eudicots

axial
nectary
common
G>5 laterally connate with abaxial nectaries; fr aggregate of follicles Trochodendrales* Trochodendraceae
mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic;
Buxales* Buxaceae (incl. Didymelaceae) Haptanthaceae
benzylisoquinolines absent

flw tepals ± uniform or missing; pregnan pseudoalkaloids

unisexual, dioecious; lvs toothed, sec. veins palmate;

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

Celastraceae Lepidobotryaceae
seeds often with aril (red-orange);
infl cymose Celastrales (incl. Hippocrateaceae, Brexiaceae) Parnassiaceae
Achariaceae Euphorbiaceae Ochnaceae Podostemaceae
lvs margins toothed;
flw G often tricarpellate Malpighiales Chrysobalanaceae
Clusiaceae
Hypericaceae
Linaceae
Passifloraceae
Phyllanthaceae
Rhizophoraceae
Salicaceae
Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae
Eurosids I

lvs often compound, pulvini (sleep movement);

flw A5 or multiple, branched style common;
mucilage cells; oxalates Oxalidales Brunelliaceae
Cephalotaceae
Connaraceae
Cunoniaceae
Elaeocarpaceae
Huaceae
Oxalidaceae
flw often “papilionaceous”: wing, standard, keel, mostly G1,

Fabales
mostly A10; fr a pod; symbiosis with root nodule bacteria;
diverse alkaloids, NP amino acids, lectins (in Fabaceae) Fabaceae Polygalaceae Quillajaceae Surianaceae
Core Eudicots

lvs mostly simple with stipules; Barbeyaceae Elaeagnaceae Rosaceae

Rosales
flw K valvate (and hypanthium) persisting,
N fix
carpels with 1 ovule, stigma dry; dihydroflavonols Cannabaceae Moraceae Ulmaceae
embryo large, Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae)
lvs with endosperm scanty
stipules
lvs mostly alternate;
Anisophyllaceae Coriariaceae Cucurbitaceae
Cucurbitales
flw often unisexual, G mostly inferior,
parietal placentation; cucurbitacins
Begoniaceae Corynocarpaceae Datiscaceae
Rosids

lvs undivided; flw small, unisexual,

anemophilous, thus T reduced or lacking, G mostly inferior;
Betulaceae Fagaceae Myricaceae Rhoipteleaceae
infl spikes or catkins; fr 1-seeded, mostly nuts;
ectomycorrhiza; tannins, dihydroflavonols Fagales Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae
lvs with glandular teeth;
flw pentacyclic,
parts alternating,
flw A obdiplostemonous, nectary on filament;
stems jointed at nodes; ethereal oils, ellagic acid Geraniales Francoaceae Geraniaceae Ledocarpaceae Melianthaceae
G connate,
K + C (free),
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)
Myrtales
(A polyandrous),
flw K valvate, persisting, A incurved in bud, ovary inferior; ovules many;
pollen tricolporate;
trihydroxyl-flavonoids endosperm scanty; scaly bark; flavonols, myricetin Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae
woody;
Crossosomataceae Stachyuraceae
K + C, stylodia free;
hypanthium, nectary disk Crossosomatales Geissolomataceae Staphyleaceae
Strasburgeriaceae
trees; lvs spiral; extrafloral nectaries;

Picramniales*
Eurosids II

staminate flw: A = and opposite C;

bark bitter, anthraquinones Picramniaceae
mostly woody (silica/silicified);
Anacardiaceae Meliaceae Rutaceae
lvs alternate, odd-pinnately compound;
flw often imperfect, intrastaminal disk; ethereal oils Sapindales Burseraceae Nitrariaceae Sapindaceae
Simaroubaceae

vessel elements: scalariform perforations; mucilage cells;
embryo
large
lvs margins toothed, stipules cauline;
flw small, A = and opposite K, ovules 1-2/carpel; Huerteales* Dipentodontaceae Gerrardinaceae Tapisciaceae
endo- bark fibrous; hairs often stellate; Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
sperm flw K often valvate,
scanty petals contorted, A often ∞; mucilage,
cyclopropenoid fatty acids, flavones
Malvales Cistaceae Cytinaceae
Dipterocarpaceae
Muntingiaceae
Neuradaceae
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; lvs stomata cyclocytic, Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae
petiole bundles annular; fr fleshy;
calcium oxalate as crystals Berberidopsidales* Aextoxicaceae Berberidopsidaceae
woody; parasites or semiparasites; without mycorrhiza;
lvs margin entire; flw A epipetalous, Balanophoraceae Misodendraceae Opiliaceae
perianth often simple, valvate, persisting;
polyacetylenes, triterpene sapogenins, silicic acid
Santalales Loranthaceae Olacaceae Santalaceae (incl. Viscaceae)
mostly herbaceous; without mycorrhiza; Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae
G often unilocular with central placentation, Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae
pollen copate, surface spiny;
betalains or anthocyanins (latter, e.g., in Caryophyllaceae) Caryophyllales [incl. Chenopodiaceae] Droseraceae
Basellaceae Drosophyllaceae
Nyctaginaceae
Phytolaccaceae
Simmondsiaceae
Talinaceae
mostly woody; lvs mostly undivided, hydathode teeth; Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae
flw often 4-merous, K much smaller than C, persisting,
intrastaminal disk, G inferior; fr drupaceous;
diverse iridoids Cornales Cornaceae
Curtisiaceae
Grubbiaceae
Hydrangeaceae
Loasaceae
Nyssaceae
lvs teeth, theoid; nodes unilacunar; Actinidiaceae Ericaceae Polemoniaceae Sarraceniaceae
flw 5-merous, pentacyclic;
nonhydrolyzable tannins, ellagic acid, hydroquinones Ericales Balsaminaceae
Clethraceae
Fouquieriaceae
Lecythidaceae
Primulaceae
Roridulaceae
Styracaceae
Theaceae
woody; dioecious; Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae
flw small, C valvate, G unilocular; fruit indehiscent;
iridoids (aucubin), gutta
Garryales Eucommiaceae Garryaceae (incl. Aucubaceae)
Asterids

lvs opposite, colleters;

Apocynaceae (incl. Asclepiadaceae) Loganiaceae
Gentianales
flw corolla convolute in bud;
late indole alkaloids; iridoids Gentianaceae Gelsemiaceae Rubiaceae
© Theodor C.H. Cole (Heidelberg) and Hartmut H. Hilger (Berlin) 2010

sympetaly
lvs opposite; nodes 1:1; flw mostly monosymmetric, Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae
A often 2(+2); ethereal oils in gland-headed hairs;
Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae
Euasterids I

6-oxygenated flavones, rosmarinic acid,

oligosaccharides: cornoside, verbascoside (acetoside) Lamiales Byblidaceae Martyniaceae Pedaliaceae Stilbaceae
mostly sympetalous, lvs spiral, simple; nodes unilacunar;
Gesneriaceae Hydrostachyaceae Oleaceae Phrymaceae Verbenaceae
A=C,

Solanales
nectary gynoecial,
ovules unitegmic,
epipetalous
polyandry
flw petals plicate; K persisting;
diverse alkaloids, no iridoids
Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae)
endosperm cellular; rare,
G(2)
Hydroleaceae Montiniaceae Sphenocleaceae
iridoids common lvs roughly hairy; nodes unilacunar;
Boraginaceae (incl. Codon) Ehretiaceae (incl. Lennoaceae) Hydrophyllaceae
infl
cymose
ellagic acid
infl scorpioid; mostly 4 ovules;
isokestose, higher inulins, pyrrolizidine alkaloids Boraginales Cordiaceae Heliotropiaceae Wellstediaceae ?
woody; lvs serrate;

Aquifoliales
lacking
flw 1-2 ovules/carpel, C± free, K slightly connate;
fr drupe with broad stigma
Aquifoliaceae Cardiopteridaceae Stemonuraceae
early
sympetaly
fl small
flw often monosymmetric, Asteraceae Goodeniaceae Pentaphragmataceae
plunger pollination (long style);
embryo
short
nodes trilacunar;
inulin, sesquiterpenes, secoiridoids
Asterales Calyceraceae Menyanthaceae
Campanulaceae (incl. Lobeliaceae)
Rousseaceae
Stylidiaceae
mostly woody; infl racemose, C free,
Escalloniales* Escalloniaceae
Euasterids II

anthers basifixed, nectary disc

Bruniales*
woody, evergreen; nodes 1:1;
flw polysymmetric, anthers basifixed Bruniaceae Columelliaceae (incl. Desfontainiaceae)
G inferior

woody (except Apiaceae);

lvs often divided; nodes usu. multilacunar; Apiaceae Griseliniaceae Pennantiaceae
________________________________________________________________________________________
- hypothetical tree based on molecular phylogenetic data (Jan 2010); branch lengths deliberate, not expressing actual time scale;
infl mostly umbel; drupe or schizocarp
(Apiaceae: mericarp/carpophore)
A piales Araliaceae Myodocarpaceae Pittosporaceae
position of many characters on tree unclear; if a character is marked as being a potential synapomorphy at a node/for a clade,
1–few-seeded

this does not mean that all members of that clade possess that character; * orders added as of APG III (2009) woody; infl racemose, flw 4-merous,
Paracryphiales*
fr usually

- this poster depicts only the largest and most important of the currently accepted approx. 450 families (according to APweb 2010); filaments stout, capsule septicidal Paracryphiaceae
for family characteristics see: Kubitzki K, ed. (1990 ff).
lvs opposite, often basally connate;
- References: APG III (2009); Judd W et al. (2007); Simpson M (2005); Soltis DE et al. (2005);
nodes 3:3; buds with scales; Adoxaceae Diervillaceae Linnaeaceae
Dipsacales
Stevens PF (2010) APweb – www.mobot.org/MOBOT/research/APweb; Watson/Dallwitz (2009) delta-intkey.com/angio/
flw often monosymmetric; Valerianaceae
- Thanks to Christoph Dobeš, Ray F. Evert, Peter H. Raven, Douglas E. Soltis, Peter F. Stevens, Maximilian Weigend, Michael Wink
K persistent in fruit; secoiridoids Caprifoliaceae Dipsacaceae Morinaceae