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) Amborellaceae A ∞ AMBORELL A LES * G aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A3– ; ∞ embryo sac 4-nucleate; N R seeds operculate, perisperm; mucilage; alkaloids (no benzylisoquinolines) NYMPH A E A LES * Cabombaceae Hydatellaceae Nymphaeaceae I A woody, vessels solitary; flw T>10, ∞A , G ca.9; embryo sac 4-nuceate; T D tiglic acid, aromatic terpenoids AUSTROB A ILEY A LES Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae E lvs opposite, interpetiolar stipules; nodes swollen; A flw small T0–3, A1–5, G1, 1 apical ovule/carpel CHLOR A NTH A LES * Chloranthaceae woody; pollen uniporate; aromatic terpenoids CA NELL A LES Canellaceae Winteraceae nodes trilacunar ± herbaceous; lvs two-ranked, leaf base sheathing; Aristolochiaceae Piperaceae single adaxial prophyll; swollen nodes PIPER A LES Hydnoraceae Saururaceae sesquiterpenes (pellucid dots) (pellucid ethereal oils in spherical idioblasts idioblasts spherical in oils ethereal woody; lvs opposite; flw with hypanthium, staminodes frequent, Calycanthaceae Hernandiaceae Monimiaceae often valvate anthers; carpels with 1 ovule; embryo large IOSPERMS LA UR A LES Gomortegaceae Lauraceae Siparunaceae G Mag NOLIIDS woody; pith septate; lvs two-ranked; ovules with obturator; N Annonaceae Eupomatiaceae Magnoliaceae features as in endosperm ruminate Mag NOLI A LES Degeneriaceae Himantandraceae Myristicaceae A “Early Angiosperms” infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules; ovules atropous, with epidermal perisperm and copious endosperm; idioblasts with ethereal oils Acoraceae RLY ACOR A LES A mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules; Alismataceae (incl. Limnocharitaceae) Juncaginaceae Ruppiaceae embryo sac 8-nucleate sac embryo benzylisoquinoline alkaloids; alkaloids; benzylisoquinoline E flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green ALISM A T A LES Aponogetonaceae Butomaceae Posidoniaceae Scheuchzeriaceae woody; vessels absent, Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae eustele; nodes; stem with ring of bundles; sieve tube plastids Ca oxalate fr a follicle; East Asia PETROS A VI A LES * Petrosaviaceae with starch grains; MONOCOTS raphides lvs simple, persistent, entire; endosperm often twining vines; lvs often reticulate; flw parts free, strobilar, atactostele (scattered bundles), nuclear Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae perfect, P parts varying, no secondary thickening; helobial, ovary often inferior, style short, branched; steroid sapogenins/alkaloids DIOSCORE A LES often in threes, mostly herbaceous; nectaries when present weakly differentiated, pollen monosulcate; some woody (with terminally tufted leaves), septal Cyclanthaceae Pandanaceae Velloziaceae stamen with broad filaments, sieve tube plastids with flw spadix PA ND A N A LES pollen monosulcate; protein crystals; often geophytes (bulbs, tubers, rhizomes); G apocarpous sympodial branching; Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae (style short in most), vascular bundles in stem scattered; flw tepals sometimes spotted, nectaries at tepals; many seeds, seeds coat (testa) cellular; nectaries absent; lvs parallel-veined, entire, phytomelan lacking; fructans in stems, chelidonic acid, steroid saponins LILI A LES 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) filaments narrow, flw nonspotted tepals, septal nectaries; capsule or berry; Asparagaceae (incl. Agavaceae, Hyacinthaceae, Ruscaceae) anthers broadly attached, seed coat obliterated or with phytomelan SP A R aga LES septal nectary; single cotyledon; A Hypoxidaceae Iridaceae Lanariaceae Orchidaceae adventitious roots monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate; Xanthorrhoeaceae (incl. Asphodelaceae, Hemerocallidaceae) intense primary growth, large apical meristem; infl with spathe;alkaloids AREC A LES Arecaceae mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent; Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae lvs grassy; flw often anemophilous, minute, chaffy, without nectaries PO A LES Cyperaceae Juncaceae Rapateaceae Typhaceae (incl. Sparganiaceae) flw often irregular; few fertile stamens; COMMELINIDS infl thyrsus of scorpioid cymes; phenylphenalenones COMMELIN A LES 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; G inferior, A often strongly modified/reduced; Cannaceae Heliconiaceae Marantaceae Strelitziaceae cuticular waxes often in rodlets seeds arillate; silicic acid aggregated into scallops ZIN G IBER A LES 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 CER A TOPHYLL A LES Ceratophyllaceae lvs often divided; flw whorled, P single or multiple whorls, G apocarpous/paracarpous, superior; Berberidaceae Eupteleaceae Menispermaceae Ranunculaceae berberines RA NUNCUL A LES Circaeasteraceae Lardizabalaceae Papaveraceae woody; flw 4–5-merous, nectary: disk, A epipetalous, G syncarpous; pollen colporate; endosperm helobial; triterpenoids SA BI A LES Sabiaceae mostly woody; flw tepals often 4-merous, A epitepalous, connectives sometimes with apical appendage PROTE A LES Nelumbonaceae Platanaceae Proteaceae axial woody; vessels lacking; idioblasts in cortex; flw tepals missing,A ∞, nectary G>5 laterally connate with abaxial nectaries; fr aggregate of follicles common TROCHODENDR A LES * Trochodendraceae mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic; flw tepals ± uniform or missing;pregnan pseudoalkaloids BUX A LES * Buxaceae (incl. Didymelaceae) Haptanthaceae UDICOTS unisexual, dioecious; lvs toothed, sec. veins palmate; flw tepals small to lacking; E ellagic acid GUNNER A LES Gunneraceae Myrothamnaceae pollen tricolpate; mostly woody; lvs veins proceed to apex & teeth; flw K5–∞, persisting; flw K/C/P opp A; mostly A∞, G mostly slightly connate; filaments rather narrow, seeds often with aril; fr often both ventrally/dorsally dehiscent Dilleniaceae anthers basifixed; DILLENI A LES nodes trilacunar; lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent; benzylisoquinolines absent benzylisoquinolines stomata anomocytic; fr mostly dry, dehiscent; Altingiaceae Daphniphyllaceae Hamamelidaceae ethereal oils absent myricetin, flavonols SA XIFR aga LES Cercidiphyllaceae Grossulariaceae Paeoniaceae often tendrillar vines; lvs often divided and with glandular teeth; Crassulaceae Haloragaceae Saxifragaceae A epipetalous, 2 ovules per carpel; raphides, pearl glands; berries VIT A LES * Vitaceae cork origin deep-seated; endosperm lacking; resinous, lignans/neolignans, harman alkaloids ZY G OPHYLL A LES * Krameriaceae Zygophyllaceae flw small, G often 3-merous, nectary: intrastaminal disk; seeds often with aril (red-orange); Celastraceae Lepidobotryaceae infl cymose CEL A STR A LES (incl. Hippocrateaceae, Brexiaceae) Parnassiaceae Achariaceae Euphorbiaceae Ochnaceae Podostemaceae ellagic acid tannins lvs margins toothed; Chrysobalanaceae Hypericaceae Passifloraceae Rhizophoraceae flw G often tricarpellate MA LPI G HI A LES Clusiaceae Linaceae Phyllanthaceae Salicaceae Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae I lvs often compound, pulvini (sleep movement); flw A5 or multiple, branched style common; Brunelliaceae Connaraceae Elaeocarpaceae mucilage cells; oxalates X A LID A LES Oxalidaceae O Cephalotaceae Cunoniaceae Huaceae 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) Fabaceae Polygalaceae Quillajaceae Surianaceae UROSIDS FA B A LES E lvs mostly simple with stipules; Barbeyaceae Elaeagnaceae Rosaceae flw K valvate (and hypanthium) persisting, N fix carpels with 1 ovule, stigma dry; dihydroflavonols ROS A LES Cannabaceae Moraceae Ulmaceae embryo large, Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae) lvs with UDICOTS endosperm scanty stipules lvs mostly alternate; E flw often unisexual, G mostly inferior, Anisophyllaceae Coriariaceae Cucurbitaceae parietal placentation; cucurbitacins CUCURBIT A LES Begoniaceae Corynocarpaceae Datiscaceae lvs undivided; flw small, unisexual, anemophilous, thus T reduced or lacking, G mostly inferior; ORE infl spikes or catkins; fr 1-seeded, mostly nuts; Betulaceae Fagaceae Myricaceae Rhoipteleaceae C ectomycorrhiza; tannins, dihydroflavonols aga LES OSIDS F Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae lvs with glandular teeth; R flw pentacyclic, flw A obdiplostemonous, nectary on filament; Francoaceae Geraniaceae Ledocarpaceae Melianthaceae parts alternating, stems jointed at nodes; ethereal oils, ellagic acid GER A NI A LES G connate, lvs opposite, colleters (glandular hair on adaxial surface of petiole base), Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae) K + C (free), stipules small (if any); cork deep seated;