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 ∞ Am b o r e l l a l e s * g aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A3– ; ∞ embryo sac 4-nuceate; N r seeds operculate, perisperm; mucilage; alkaloids (no benzylisoquinolines) Ny m p h a e a l e s * 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 Au s t r o b a i l e y a l e s Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae e lvs opposite, interpetiolar stipules; nodes swollen; A flw small T0–3, A1–5, G1, 1 apical ovule/carpel; sesquiterpenes Ch l o r a n t h a l e s * Chloranthaceae woody; pollen uniporate; aromatic terpenoids Ca n e l l a l e s Canellaceae Winteraceae nodes trilacunar ± herbaceous; lvs two-ranked, leaf base sheathing; Aristolochiaceae Piperaceae

sesquiterpenes single adaxial prophyll; swollen nodes; sesquiterpenes Pi p e r a l e s Hydnoraceae Saururaceae

(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

i o s p e r m s La u r a l e s Gomortegaceae Lauraceae Siparunaceae g Mag n o l i i d s woody; pith septate; lvs two-ranked; ovules with obturator; n Annonaceae Eupomatiaceae Magnoliaceae features as in endosperm ruminate; sesquiterpenes Mag n o l i a l e s 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

r l y Ac o r a l e s

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 Al i s m a t a l e s 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 Pe t r o s a v i a l e s * Petrosaviaceae with starch grains; Mo n o c o t s 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 Di o s c o r e a l e s 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 n d a n a l e s 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 Li l i a l e s 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 s p a r aga l e s 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 Ar e c a l e s Arecaceae mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent; Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae lvs grassy; flw often anemophilous, minute, chaffy, without nectaries Po a l e s Cyperaceae Juncaceae Rapateaceae Typhaceae (incl. Sparganiaceae) flw often irregular; few fertile stamens; Co m m e l i n i d s infl thyrsus of scorpioid cymes; phenylphenalenones Co m m e l i n a l e s 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 Zi n g i b e r a l e s 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 Ce r a t o p h y l l a l e s Ceratophyllaceae lvs often divided; flw whorled, P single or multiple whorls, G apocarpous/paracarpous, superior; Berberidaceae Eupteleaceae Menispermaceae Ranunculaceae berberines Ra n u n c u l a l e s Circaeasteraceae Lardizabalaceae Papaveraceae woody; flw 4–5-merous, nectary: disk, A epipetalous, G syncarpous; pollen colporate; endosperm helobial; triterpenoids Sa b i a l e s Sabiaceae mostly woody; flw tepals often 4-merous, A epitepalous, connectives sometimes with apical appendage Pr o t e a l e s 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 Tr o c h o d e n d r a l e s * Trochodendraceae mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic; flw tepals ± uniform or missing;pregnan pseudoalkaloids Bu x a l e s * Buxaceae (incl. Didymelaceae) Haptanthaceae

u d i c o t s unisexual, dioecious; lvs toothed, sec. veins palmate; flw tepals small to lacking; E ellagic acid Gu n n e r a l e s 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; Di l l e n i a l e s 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 x i f r aga l e s 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 Vi t a l e s * Vitaceae cork origin deep-seated; endosperm lacking; resinous, lignans/neolignans, harman alkaloids Zy g o p h y l l a l e s * Krameriaceae Zygophyllaceae flw small, G often 3-merous, nectary: intrastaminal disk; seeds often with aril (red-orange); Celastraceae Lepidobotryaceae infl cymose Ce l a s t r a l e s (incl. Hippocrateaceae, Brexiaceae) Parnassiaceae Achariaceae Euphorbiaceae Ochnaceae Podostemaceae ellagic acid tannins lvs margins toothed; Chrysobalanaceae Hypericaceae Passifloraceae Rhizophoraceae flw G often tricarpellate Ma l p i g h i a l e s 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 l i d a l e s 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 u r o s i d s Fa b a l e s

E lvs mostly simple with stipules; Barbeyaceae Elaeagnaceae Rosaceae flw K valvate (and hypanthium) persisting, N fix carpels with 1 ovule, stigma dry; dihydroflavonols Ro s a l e s Cannabaceae Moraceae Ulmaceae embryo large, Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae) lvs with

u d i c o t s endosperm scanty stipules lvs mostly alternate; flw often unisexual, G mostly inferior, Anisophyllaceae Coriariaceae Cucurbitaceae E parietal placentation; cucurbitacins Cu c u r b i t a l e s Begoniaceae Corynocarpaceae Datiscaceae lvs undivided; flw small, unisexual, anemophilous, thus T reduced or lacking, G mostly inferior; o r e infl spikes or catkins; fr 1-seeded, mostly nuts; Betulaceae Fagaceae Myricaceae Rhoipteleaceae C ectomycorrhiza; tannins, dihydroflavonols aga l e s o s i d s F Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae lvs with glandular teeth;

R flw pentacyclic, flw A obdiplostemonous, nectary on filament; Francoaceae Melianthaceae Ledocarpaceae Geraniaceae parts alternating, stems jointed at nodes; ethereal oils, ellagic acid Ge r a n i a l e s 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; (A polyandrous), flw K valvate, persisting, A incurved in bud, ovary inferior; ovules many; Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae) pollen tricolporate; trihydroxyl-flavonoids endosperm scanty; scaly bark; flavonols, myricetin My r t a l e s Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae woody; K + C, stylodia free; Crossosomataceae Stachyuraceae Strasburgeriaceae hypanthium, nectary disk Cr o s s o s o m a t a l e s Geissolomataceae Staphyleaceae trees; lvs spiral; extrafloral nectaries;

staminate flw: A = and opposite C; II

bark bitter, anthraquinones Pi c r a m n i a l e s * Picramniaceae mostly woody (silica/silicified); Anacardiaceae Meliaceae Rutaceae lvs alternate, odd-pinnately compound; Simaroubaceae flw often imperfect, intrastaminal disk;ethereal oils Sa p i n d a l e s Burseraceae Nitrariaceae Sapindaceae vessel elements: scalariform perforations; mucilage cells; lvs margins toothed, stipules cauline; u r o s i d s embryo Dipentodontaceae Gerrardinaceae Tapisciaceae

large flw small, A = and opposite K, ovules 1-2/carpel; Hu e r t e a l e s * E endo- bark fibrous; hairs often stellate; Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae) sperm flw K often valvate, scanty petals contorted, A often ∞; mucilage, Cistaceae Cytinaceae Muntingiaceae Sarcolaenaceae cyclopropenoid fatty acids, flavones Ma l v a l e s Dipterocarpaceae Neuradaceae 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 Br a s s i c a l e s Brassicaceae Cleomaceae Moringaceae Tovariaceae woody; lvs stomata cyclocytic, Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae petiole bundles annular; fr fleshy; calcium oxalate as crystals Berberidopsid a l e s * Aextoxicaceae Berberidopsidaceae woody; parasites or semiparasites; without mycorrhiza; lvs margin entire; flw A epipetalous, Balanophoraceae Misodendraceae Opiliaceae perianth often simple, valvate, persisting; Loranthaceae Olacaceae Santalaceae (incl. Viscaceae) polyacetylenes, triterpene sapogenins, silicic acid Sa n t a l a l e s mostly herbaceous; without mycorrhiza; Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae G often unilocular with central placentation, Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae pollen copate, surface spiny; [incl. Chenopodiaceae] Droseraceae Nyctaginaceae Simmondsiaceae betalains or anthocyanins (latter, e.g., in Caryophyllaceae) Ca r y o p h y l l a l e s Basellaceae Drosophyllaceae Phytolaccaceae 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; Cornaceae Grubbiaceae Loasaceae diverse iridoids o r n a l e s C Curtisiaceae Hydrangeaceae Nyssaceae lvs teeth, theoid; nodes unilacunar; Actinidiaceae Ericaceae Polemoniaceae Sarraceniaceae flw 5-merous, pentacyclic; nonhydrolyzable tannins, ellagic acid, hydroquinones r i c a l e s Balsaminaceae Fouquieriaceae Primulaceae Styracaceae E Clethraceae Lecythidaceae Roridulaceae Theaceae woody; dioecious; Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae flw small, C valvate, G unilocular; fruit indehiscent; iridoids (aucubin), gutta Ga r r y a l e s Eucommiaceae Garryaceae (incl. Aucubaceae) lvs opposite, colleters; flw corolla convolute in bud; Apocynaceae (incl. Asclepiadaceae) Loganiaceae late indole alkaloids; iridoids sympetaly Ge n t i a n a l e s Gentianaceae Gelsemiaceae Rubiaceae lvs opposite; nodes 1:1; flw mostly monosymmetric, Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae A often 2(+2); ethereal oils in gland-headed hairs;

s t e r i d s I 6-oxygenated flavones, rosmarinic acid, Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae

A oligosaccharides: cornoside, verbascoside (acetoside) a m i a l e s

L Byblidaceae Martyniaceae Pedaliaceae Stilbaceae Gesneriaceae Hydrostachyaceae Oleaceae Phrymaceae Verbenaceae mostly sympetalous, A=C, lvs spiral, simple; nodes unilacunar; nectary gynoecial, epipetalous flw petals plicate; K persisting; Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae) ovules unitegmic, polyandry diverse alkaloids, no iridoids rare, So l a n a l e s endosperm cellular; s t e r i d s Hydroleaceae Montiniaceae Sphenocleaceae iridoids common G(2) a lvs roughly hairy; nodes unilacunar; infl u infl scorpioid; mostly 4 ovules; Boraginaceae (incl. Codon) Ehretiaceae (incl. Lennoaceae) Hydrophyllaceae cymose isokestose, higher inulins, pyrrolizidine alkaloids

E o r ag i n a l e s ellagic acid B Cordiaceae Heliotropiaceae Wellstediaceae ? woody; lvs serrate; lacking flw 1-2 ovules/carpel, C± free, K slightly connate; Aquifoliaceae Cardiopteridaceae Stemonuraceae early fr drupe with broad stigma Aq u i f o l i a l e s sympetaly flw often monosymmetric, Asteraceae Goodeniaceae Pentaphragmataceae fl small plunger pollination (long style); embryo nodes trilacunar; Calyceraceae Menyanthaceae Rousseaceae short As t e r a l e s inulin, sesquiterpenes, secoiridoids Campanulaceae (incl. Lobeliaceae) Stylidiaceae mostly woody; infl racemose, C free,

anthers basifixed, nectary disc s c a l l o n i a l e s * Escalloniaceae II E woody, evergreen; nodes 1:1;

Theodor C.H. Cole (Heidelberg) and Hartmut H. Hilger (Berlin) 2010 (Berlin) Hilger H. Hartmut and (Heidelberg) Cole C.H. Theodor flw polysymmetric, anthers basifixed Bruniaceae Columelliaceae (incl. Desfontainiaceae)

© © Br u n i a l e s * woody (except Apiaceae);

______lvs often divided; nodes usu. multilacunar; Apiaceae Griseliniaceae Pennantiaceae G inferior infl mostly umbel; drupe or schizocarp

- hypothetical tree based on molecular phylogenetic data (Jan 2010); branch lengths deliberate, not expressing actual time scale; s t e r i d s (Apiaceae: mericarp/carpophore) Ap i a l e s 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, a 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, u - this poster depicts only the largest and most important of the currently accepted approx. 450 families (according to APweb 2010); filaments stout, capsule septicidal Pa r a c r y p h i a l e s * Paracryphiaceae for family characteristics see: Kubitzki K, ed. (1990 ff). References: APG III (2009); Judd W et al. (2007); Simpson M (2005); E lvs opposite, often basally connate; Soltis DE et al. (2005); Stevens PF (2010) APweb – www.mobot.org/MOBOT/research/APWeb/; Watson/Dallwitz (2009) delta-intkey. fr usually 1–few-seeded nodes 3:3; buds with scales; com/angio/ 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; Valerianaceae K persistent in fruit; secoiridoids Di p s a c a l e s Caprifoliaceae Dipsacaceae Morinaceae