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 1/1/1 MBORELLALES G A A aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A1– , embryo sac 4-nucleate R ∞ seeds operculate with perisperm but endosperm reduced or small; mucilage; alkaloids (no benzylisoquinolines) Cabombaceae Hydatellaceae Nymphaeaceae 3/6/74 NYMPHAEALES N A D woody, vessels solitary; flw T>10, ∞A , G ca.9, embryo sac 4-nucleate tiglic acid, aromatic terpenoids (incl. Illiciaceae) 3/5/100 Austrobaileyaceae Schisandraceae Trimeniaceae E AUSTROBAILEYALES A lvs opposite, interpetiolar stipules; nodes swollen A flw small T0–3, A1–5, G1, 1 apical ovule/carpel Chloranthaceae 1/4/75 CHLORANTHALES E N A woody; foliar sclereids, K and C distinct G aromatic terpenoids Canellaceae Winteraceae 2/10/125 CANELLALES R idioblasts spherical in I nodes trilacunar L ± herbaceous; lvs two-ranked, leaf base sheathing O single adaxial prophyll; swollen nodes Aristolochiaceae (incl. Hydnoraceae) Piperaceae Saururaceae 4/17/4170 IPERALES P Y sesquiterpenes S woody; lvs opposite; flw with hypanthium, staminodes frequent Calycanthaceae Hernandiaceae Monimiaceae tension wood + wood tension (pellucid dots) (pellucid ethereal oils ethereal often valvate anthers; carpels with 1 ovule; embryo large P 7/91/2858 AURALES Gomortegaceae Lauraceae Siparunaceae L AGNOLIIDS E M woody; pith septate; lvs two-ranked; ovules with obturator Annonaceae Eupomatiaceae Magnoliaceae features as in endosperm ruminate R 6/128/3140 MAGNOLIALES “Early Angiosperms” Degeneriaceae Himantandraceae Myristicaceae M infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules ovules atropous, seeds with epidermal perisperm and copious endosperm; ethereal oils Acoraceae S 1/1/2-4 ACORALES woody; vessels absent mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules Alismataceae Butomaceae Posidoniaceae Scheuchzeriaceae eustele infl ± scapose; flw G apocarpous; placentation often laminar; endosperm helobial; embryo large/green Aponogetonaceae Hydrocharitaceae Potamogetonaceae Tofieldiaceae 14/166/4660 ALISMATALES sieve tube plastids Araceae Juncaginaceae Ruppiaceae Zosteraceae with starch grains stem with ring of bundles endosperm triploid endosperm embryo sac 8-nucleate sac embryo lvs simple, persistent, entire alkaloids benzylisoquinoline fr a follicle Ca oxalate East Asia flw strobilar, perfect, parts free 1/2/3 Petrosaviaceae MONOCOTS raphides PETROSAVIALES P parts varying, often in threes, weakly differentiated scattered bundles in stem endosperm often twining vines; lvs often reticulate nuclear anthers tetrasporangiate no secondary thickening ovary often inferior, style short, branched; steroidal sapogenins/alkaloids IOSCOREALES Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae Thismiaceae helobial 5/21/1050 D stamen with broad filaments mostly herbaceous pollen monosulcate pollen monosulcate some woody (with terminally tufted leaves) G apocarpous (style short in most) sieve tube plastids with infl sometimes spathe + spadix Cyclanthaceae Pandanaceae Triuridaceae Velloziaceae 5/36/1345 ANDANALES compitum (if present) extragynoecial protein crystals P nectaries absent sympodial branching often geophytes (bulbs, tubers, rhizomes); leaf bases often not sheathing Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae siphonogamy lvs parallel-veined, entire flw tepals sometimes spotted, nectaries on tepals, anthers extrorse double fertilization > endosperm no glandular teeth many seeds; phytomelan lacking; fructans in stems, chelidonic acid ILIALES 10/67/1558 L Colchicaceae Liliaceae Petermanniaceae Smilacaceae embryo very small flw pentacyclic P 3-merous, A opp. P often geophytes incl. Agapanthaceae, Alliaceae filaments narrow capsule or berry Amaryllidaceae ( ) Hypoxidaceae Iridaceae anthers broadly attached seed coat obliterated or with phytomelan Asparagaceae (incl. Agavaceae, Hyacinthaceae, Ruscaceae) Lanariaceae Orchidaceae septal nectary 14/1122/36205 ASPARAGALES single cotyledon Tecophilaeaceae Asphodelaceae (incl. Xanthorrhoeaceae, Hemerocallidaceae) woody, often monopodial extrafloral nectaries extrafloral radicle not persistent stem-borne roots numerous lvs often palmately or pinnately pseudocompound, reduplicate-plicate intense primary growth, large apical meristem, infl often with spathe; alkaloids 1/188/2585 Arecaceae Dasypogonaceae ARECALES mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae lvs grassy; flw often anemophilous, minute, chaffy, without nectaries 15/997/18875 POALES Cyperaceae Juncaceae Rapateaceae Typhaceae (incl. Sparganiaceae) flw monosymmetric or not, few fertile stamens infl thyrsus of scorpioid cymes Commelinaceae Haemodoraceae Hanguanaceae COMMELINIDS phenylphenalenones 5/68/812 COMMELINALES Philydraceae Pontederiaceae UV-fluorescing cell walls rhizomatous, large-leafed herbs; pseudostem common (ferulic/coumaric acids) flw irregular/monosymmetric, septal nectaries silicic acid in leaves A often strongly modified/reduced, G inferior Cannaceae Heliconiaceae Marantaceae Strelitziaceae cuticular waxes often in rodlets seeds often arillate, silicic acid 8/92/~2500 aggregated into scallops ZINGIBERALES Costaceae Lowiaceae Musaceae Zingiberaceae aquatic; herbaceous; monoecious; lvs whorled, no pellucid dots; vessels lacking flw T0 or –9 10*, A1, G1, 1 apical ovule/carpel Ceratophyllaceae pollen inaperturate, pollen tube branched, hydrophilous 1/1/2 CERATOPHYLLALES lvs often divided; flw parts whorled, P single or multiple whorls Berberidaceae Eupteleaceae Menispermaceae ethereal G apocarpous/paracarpous, superior; berberines Ranunculaceae 7/199/4510 ANUNCULALES oils R Circaeasteraceae Lardizabalaceae Papaveraceae not in mostly woody; flw tepals often 4-merous idioblasts A epitepalous, connectives sometimes with apical appendage Nelumbonaceae Platanaceae Proteaceae Sabiaceae 4/85/1750 PROTEALES woody; vessels lacking; flw tepals missing, ∞A G>5 laterally connate with abaxial nectaries; fr aggregate of follicles Trochodendraceae recepta- 1/2/2 TROCHODENDRALES cular mostly woody; mostly monoecious, flw unisexual; lvs evergreen, stomata cyclocytic E nectary common flw tepals ± uniform or missing;pregnane pseudoalkaloids Buxaceae (incl. Haptanthaceae) U 1/7/120 BUXALES D dioecious, flw unisexual; lvs toothed, sec. veins palmate flw tepals small to lacking I ellagic acid Gunneraceae Myrothamnaceae 2/2/50 UNNERALES C G mostly woody; lvs if veins strong, proceed to apex of teeth absent O flw mostly K5, persisting, mostly ∞A , G mostly slightly connate seeds often with aril; fr usually follicles Dilleniaceae T 1/10/300 DILLENIALES S lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent Altingiaceae Cynomoriaceae Haloragaceae Peridiscaceae fr mostly dry, dehiscent pollen tricolpate myricetin, flavonols Cercidiphyllaceae Daphniphyllaceae Hamamelidaceae Paeoniaceae protandry common 15/112/2500 SAXIFRAGALES flw K/C/P opp A often tendrillar vines; lvs often divided and with glandular teeth Crassulaceae Grossulariaceae Iteaceae Saxifragaceae filaments rather narrow nodes 3:3 A epipetalous, 2 ovules per carpel; raphides, pearl glands benzylisoquinolines berries ae stomata anomocytic 1/14/850 ITALES Vitace microsporogenesis V simultaneous cork origin deep-seated endosperm lacking resinous, lignans/neolignans, harman alkaloids Krameriaceae Zygophyllaceae 2/24/345 ZYGOPHYLLALES stipules infl cymose, flw small G often 3-merous, nectary often intrastaminal disk seeds often arillate (red-orange) or winged Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae) Lepidobotryaceae 2/94/1355 CELASTRALES gallic acids gallic S and lvs often compound, pulvini (sleep movement) U COM clade flw A5 or multiple, branched style common Brunelliaceae Connaraceae Elaeocarpaceae P mucilage cells; oxalates XALIDALES Oxalidaceae 7/60/1845 ellagic ellagic O Cephalotaceae Cunoniaceae Huaceae E R habits and habitats extremely diverse Achariaceae Euphorbiaceae Rafflesiaceae Ochnaceae Podostemaceae F lvs margins toothed Chrysobalanaceae Hypericaceae Passifloraceae Rhizophoraceae E R A flw G often tricarpellate ALPIGHIALES C 36/716/16065 M Clusiaceae Linaceae Phyllanthaceae Salicaceae O U O B D S R I flw often “papilionaceous”: wing, standard, keel, C clawed, mostly G1 Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae R O mostly A10; fr a pod; symbiosis with root nodule bacteria I I D diverse alkaloids, NP amino acids, lectins (in Fabaceae) Fabaceae Polygalaceae Quillajaceae Surianaceae 4/754/20140 ABALES E C D S S F O S I lvs mostly simple with stipules Barbeyaceae Elaeagnaceae Rosaceae flw K valvate (and hypanthium) persisting D N fix T carpels with 1 ovule, stigma dry; dihydroflavonols OSALES Cannabaceae Moraceae Ulmaceae S 9/261/7725 R S embryo large Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae) endosperm scanty lvs mostly alternate flw often unisexual, G mostly inferior Apodanthaceae Begoniaceae Corynocarpaceae Datiscaceae parietal placentation; cucurbitacins 7/109/2935 CUCURBITALES Anisophyllaceae Coriariaceae Cucurbitaceae Tetramelaceae mostly trees; lvs mostly undivided; flw small, unisexual anemophilous, thus T reduced or lacking, G mostly inferior infl spikes or catkins; fr 1-seeded, mostly nuts Betulaceae Fagaceae Myricaceae ectomycorrhiza; tannins, dihydroflavonols Ticodendraceae 7/33/1005 FAGALES Casuarinaceae Juglandaceae