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) MBORELLALES Amborellaceae 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) NYMPHAEALES Cabombaceae Hydatellaceae Nymphaeaceae I A D woody, vessels solitary; flw T>10, ∞A , G ca.9; embryo sac 4-nucleate tiglic acid, aromatic terpenoids Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae T E AUSTROBAILEYALES A lvs opposite, interpetiolar stipules; nodes swollen E flw small T0–3, A1–5, G1, 1 apical ovule/carpel CHLORANTHALES Chloranthaceae A woody; pollen uniporate R aromatic terpenoids CANELLALES Canellaceae Winteraceae L nodes trilacunar ± herbaceous; lvs two-ranked, leaf base sheathing Y single adaxial prophyll; swollen nodes IPERALES Aristolochiaceae Hydnoraceae Piperaceae Saururaceae P sesquiterpenes woody; lvs opposite; flw with hypanthium, staminodes frequent Calycanthaceae Hernandiaceae Monimiaceae (pellucid dots) (pellucid ethereal oils in spherical idioblasts idioblasts spherical in oils ethereal A often valvate anthers; carpels with 1 ovule; embryo large LAURALES Gomortegaceae Lauraceae Siparunaceae AGNOLIIDS N M woody; pith septate; lvs two-ranked; ovules with obturator Annonaceae Eupomatiaceae Magnoliaceae G features as in endosperm ruminate MAGNOLIALES “Early Angiosperms” Degeneriaceae Himantandraceae Myristicaceae I infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules O ovules atropous, with epidermal perisperm and copious endosperm; idioblasts with ethereal oils ACORALES Acoraceae Alismataceae (incl. Limnocharitaceae) Juncaginaceae Ruppiaceae S mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green LISMATALES Aponogetonaceae Butomaceae Posidoniaceae Scheuchzeriaceae P A Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae embryo sac 8-nucleate sac embryo E alkaloids; benzylisoquinoline stem with ring of bundles Ca oxalate fr a follicle; East Asia Petrosaviaceae R MONOCOTS raphides PETROSAVIALES endosperm M atactostele (scattered bundles) often twining vines; lvs often reticulate nuclear Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae no secondary thickening helobial ovary often inferior, style short, branched; steroid sapogenins/alkaloids DIOSCOREALES S mostly herbaceous nectaries when present pollen monosulcate septal some woody (with terminally tufted leaves) woody; vessels absent sieve tube plastids with flw spadix ANDANALES Cyclanthaceae Pandanaceae Velloziaceae eustele; nodes protein crystals P sieve tube plastids sympodial branching often geophytes (bulbs, tubers, rhizomes) with starch grains vascular bundles in stem scattered flw tepals sometimes spotted, nectaries at tepals; many seeds, seeds coat (testa) cellular Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae lvs simple, persistent, entire lvs parallel-veined, entire phytomelan lacking; fructans in stems, chelidonic acid, steroid saponins LILIALES Colchicaceae Liliaceae Petermanniaceae Smilacaceae flw parts free, strobilar, no glandular teeth perfect, P parts varying, flw pentacyclic, P 3-merous, A opp. P often geophytes; anomalous sec. growth often in threes, filaments narrow capsule or berry Amaryllidaceae (incl. Agapanthaceae, Alliaceae) Hypoxidaceae Iridaceae weakly differentiated, anthers broadly attached, seed coat obliterated or with phytomelan (incl. Agavaceae, Hyacinthaceae, Ruscaceae) stamen with broad filaments septal nectary; single cotyledon ASPARAGALES Asparagaceae Lanariaceae Orchidaceae pollen monosulcate adventitious roots Tecophilaeaceae Xanthorrhoeaceae (incl. Asphodelaceae, Hemerocallidaceae) G apocarpous (style short in most) monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate nectaries absent intense primary growth, large apical meristem embryo very small infl with spathe;alkaloids Arecaceae aromatic terpenoids ARECALES 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 infl thyrsus of scorpioid cymes COMMELINIDS phenylphenalenones Commelinaceae Haemodoraceae Pontederiaceae COMMELINALES 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 INGIBERALES aggregated into scallops Z 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 Ranunculaceae RANUNCULALES Circaeasteraceae Lardizabalaceae Papaveraceae woody; flw 4–5-merous, nectary: disk, A epipetalous, G syncarpous pollen colporate; endosperm helobial; triterpenoids Sabiaceae SABIALES mostly woody; flw tepals often 4-merous A epitepalous, connectives sometimes with apical appendage PROTEALES Nelumbonaceae Platanaceae Proteaceae axial woody; vessels lacking; idioblasts in cortex; flw tepals missing, A∞ E nectary G>5 laterally connate with abaxial nectaries; fr aggregate of follicles U common TROCHODENDRALES Trochodendraceae D mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic flw tepals ± uniform or missing;pregnan pseudoalkaloids I BUXALES Buxaceae Didymelaceae C unisexual, dioecious; lvs toothed, sec. veins palmate flw tepals small to lacking O ellagic acid UNNERALES Gunneraceae Myrothamnaceae T G mostly woody; lvs veins proceed to apex & teeth; flw K5–∞, persisting S mostly A∞, G mostly slightly connate seeds often with aril; fr often both ventrally/dorsally dehiscent Dilleniaceae pollen tricolpate DILLENIALES flw K/C/P opp A lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent benzylisoquinolines absent benzylisoquinolines filaments rather narrow fr mostly dry, dehiscent Altingiaceae Daphniphyllaceae Hamamelidaceae anthers basifixed myricetin, flavonols Paeoniaceae nodes trilacunar SAXIFRAGALES Cercidiphyllaceae Grossulariaceae stomata anomocytic often tendrillar vines; lvs often divided and with glandular teeth Crassulaceae Haloragaceae Saxifragaceae ethereal oils absent A epipetalous, 2 ovules per carpel; raphides, pearl glands berries stipules VITALES Vitaceae nodes cork origin deep-seated 3:3 endosperm lacking resinous, lignans/neolignans, harman alkaloids ZYGOPHYLLALES Krameriaceae Zygophyllaceae R flw small, G often 3-merous, nectary: intrastaminal disk seeds often with aril (red-orange) O Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae) Lepidobotryaceae S infl cymose CELASTRALES I lvs often compound, pulvini (sleep movement) ellagic acid tannins Brunelliaceae Connaraceae Elaeocarpaceae D flw A5 or multiple, branched style common Oxalidaceae S mucilage cells; oxalates OXALIDALES Cephalotaceae Cunoniaceae Huaceae F Achariaceae Euphorbiaceae Rafflesiaceae Ochnaceae Podostemaceae lvs margins toothed Chrysobalanaceae Hypericaceae Passifloraceae Rhizophoraceae A flw G often tricarpellate MALPIGHIALES B Clusiaceae Linaceae Phyllanthaceae Salicaceae flw often “papilionaceous”: wing, standard, keel, mostly G1 Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae I 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 U lvs mostly simple with stipules Barbeyaceae Elaeagnaceae Rosaceae O N fix flw K valvate (and hypanthium) persisting R D carpels with 1 ovule, stigma dry; dihydroflavonols ROSALES Cannabaceae Moraceae Ulmaceae E I embryo large Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae) C endosperm scanty lvs mostly alternate flw often unisexual, G mostly inferior Anisophyllaceae Coriariaceae Cucurbitaceae O parietal placentation; cucurbitacins Tetramelaceae CUCURBITALES Begoniaceae Corynocarpaceae Datiscaceae T lvs undivided; flw small, unisexual S anemophilous, thus T reduced or lacking, G mostly inferior infl spikes or catkins; fr 1-seeded, mostly nuts Betulaceae Fagaceae Myricaceae Rhoipteleaceae flw pentacyclic ectomycorrhiza; tannins, dihydroflavonols FAGALES Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae parts alternating lvs with glandular teeth G connate flw A obdiplostemonous, nectary on filament K + C (free) Francoaceae Geraniaceae Ledocarpaceae Melianthaceae (A polyandrous) stems jointed at nodes; ethereal oils, ellagic acid GERANIALES pollen tricolporate lvs opposite, colleters (glandular hair on adaxial surface of petiole base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae) trihydroxyl-flavonoids stipules small (if any); cork deep seated flw K valvate, persisting, A incurved in bud, ovary inferior; ovules many Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae) endosperm scanty; scaly bark; flavonols, myricetin