Angiosperm Phylogeny Flowering Systematics A woody; dioecious; flw T5–8, A∞, G5–8; 1 ovule/carpel; embryo sac 9-nucleate; 1 species (New Caledonia) Amborellales Amborellaceae g N aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A3–∞; embryo sac 4-nuceate; 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 Austrobaileyales Austrobaileyaceae Illiciaceae (incl. Schisandraceae) Trimeniaceae e A lvs opposite, interpetiolar stipules; nodes swollen; flw small T0–3, A1–5, G1, 1 apical ovule/carpel; sesquiterpenes Chloranthales Chloranthaceae woody; pollen uniporate; vessels 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 woody; lvs opposite; flw with hypanthium, staminodes frequent, Calycanthaceae Hernandiaceae Monimiaceae often valvate anthers; carpels with 1 ovule; embryo large

La u r a l e s Gomortegaceae Lauraceae Siparunaceae (pellucid dots) (pellucid ethereal oils in spherical idioblasts idioblasts spherical in oils ethereal a g n o l i i d s M woody; pith septate; lvs two-ranked; ovules with obturator; Annonaceae Eupomatiaceae Magnoliaceae features as in endosperm ruminate; sesquiterpenes a g n o l i a l e s “Early Angiosperms” M Degeneriaceae Himantandraceae Myristicaceae

i o s p e r m s infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules;

g ovules atropous, with epidermal perisperm and copious endosperm; idioblasts with ethereal oils Ac o r a l e s Acoraceae n

mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules; Alismataceae Butomaceae Limnocharitaceae Ruppiaceae A

flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green Aponogetonaceae Juncaginaceae Posidoniaceae Scheuchzeriaceae embryo sac 8-nucleate sac embryo benzylisoquinoline alkaloids; alkaloids; benzylisoquinoline Al i s m a t a l e s Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae stem with ring of bundles;

a r l y Ca oxalate fr a follicle; East Asia Petrosaviales Petrosaviaceae

Mo n o c o t s as raphides E often twining vines; lvs often reticulate; atactostele (scattered bundles), Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae no secondary thickening; ovary often inferior, style short, branched; steroid sapogenins/alkaloids Dioscoreales woody; vessels absent, mostly herbaceous; some woody (with terminally tufted leaves), eustele; nodes; pollen monosulcate; Cyclanthaceae Pandanaceae Velloziaceae sieve tube plastids sieve tube plastids with flw spadix Pa n d a n a l e s with starch grains; protein crystals; often geophytes (bulbs, tubers, rhizomes); lvs simple, persistent, entire; sympodial branching; Alstroemeriaceae Corsiaceae Melanthiaceae flw parts free, strobilar, vascular bundles in stem scattered; flw tepals sometimes spotted, nectaries at tepals; many seeds, seeds coat (testa) cellular;

perfect, P parts varying, lvs parallel-veined, entire, nuclear/helobial, endosperm phytomelan lacking; fructans in stems, chelidonic acid, steroid saponins Li l i a l e s Colchicaceae Liliaceae Smilacaceae nectaries when present septal present when nectaries often in threes, no glandular teeth; weakly differentiated, flw pentacyclic, P 3-merous, A opp. P, often geophytes; anomalous sec. growth; Alliaceae (incl. Agapanthaceae, Amaryllidaceae) stamen with broad filaments, filaments narrow, flw nonspotted tepals, septal nectaries; capsule or berry; Asparagaceae (incl. Agavaceae, Hyacinthaceae, Ruscaceae) pollen monosulcate; anthers broadly attached, seed coat obliterated or with phytomelan s p a r a g a l e s G apocarpous septal nectary; single cotyledon; A Hypoxidaceae Iridaceae Lanariaceae Orchidaceae (style short in most), adventitious roots nectaries absent; monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate; Xanthorrhoeaceae (incl. Asphodelaceae, Hemerocallidaceae) embryo very small; intense primary growth, large apical meristem; aromatic terpenoids infl with spathe;alkaloids Ar e c a l e s Arecaceae mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent; Bromeliaceae Eriocaulaceae Poaceae Restionaceae Typhaceae lvs grassy; flw often anemophilous, minute, chaffy, without nectaries Po a l e s Cyperaceae Juncaceae Rapateaceae Sparganiaceae Xyridaceae flw often irregular; few fertile stamens; Co m m e l i n i d s 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; 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 Ceratophyllales Ceratophyllaceae lvs often divided; flw whorled, P single or multiple whorls, G apocarpous/paracarpous, superior; Berberidaceae Lardizabalaceae Papaveraceae berberines Ranunculales Eupteleaceae Menispermaceae Ranunculaceae 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 , u d i c o t s nectary ∞ E 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 pollen tricolpate; flw K/C/P opp A; mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic; filaments rather narrow, flw tepals ± uniform or missing;pregnan pseudoalkaloids Buxaceae Didymelaceae anthers basifixed; Bu x a l e s nodes trilacunar; unisexual, dioecious; lvs toothed, sec. veins palmate; stomata anomocytic; flw tepals small to lacking; ethereal oils absent ellagic acid Gu n n e r a l e s Gunneraceae Myrothamnaceae woody; lvs stomata cyclocytic, petiole bundles annular; fr fleshy; calcium oxalate as crystals Be r b e r i d o p s i d a l e s Aextoxicaceae Berberidopsidaceae mostly woody; lvs veins proceed to apex & teeth; flw K5– , persisting; benzylisoquinolines absent benzylisoquinolines ∞ mostly A∞, G mostly slightly connate; Dilleniaceae seeds often with aril; fr often both ventrally/dorsally dehiscent Di l l e n i a l e s mostly herbaceous; without mycorrhiza; Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae G often unilocular with central placentation, Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae THCpollen copate, surface spiny; [+Chenopodiaceae] Droseraceae Nyctaginaceae Simmondsiaceae betalains or anthocyanins (latter, e.g., in Caryophyllaceae) Caryophyllales Basellaceae Drosophyllaceae Phytolaccaceae Talinaceae woody; parasites or semiparasites; without mycorrhiza; Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae lvs margin entire; flw A epipetalous, perianth often simple, valvate, persisting; polyacetylenes, triterpene sapogenins, silicic acid Balanophoraceae Misodendraceae Opiliaceae Sa n t a l a l e s Loranthaceae Olacaceae Santalaceae (incl. Viscaceae) lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent; fr mostly dry, dehiscent; Altingiaceae Daphniphyllaceae Hamamelidaceae myricetin, flavonols ellagic acid Sa x i f r a g a l e s Cercidiphyllaceae Grossulariaceae Paeoniaceae tannins Crassulaceae Haloragaceae Saxifragaceae often tendrillar vines; lvs often divided and with glandular teeth; A epipetalous, 2 ovules per carpel; raphides, pearl glands; berries Vi t a l e s Vitaceae woody; K + C, stylodia free; hypanthium, nectary disk Crossosomatales Crossosomataceae Stachyuraceae Staphyleaceae lvs with glandular teeth; flw A obdiplostemonous, nectary on filament; stems jointed at nodes; ethereal oils, ellagic acid Ge r a n i a l e s Francoaceae Geraniaceae Melianthaceae lvs opposite, colleters (glandular hair on adaxial surface of petiole base), lvs with stipules small (if any); cork deep seated; Combretaceae Myrtaceae Onagraceae stipules flw K valvate, persisting, A incurved in bud, ovary inferior; ovules many; endosperm scanty; scaly bark; flavonols, myricetin My r t a l e s Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae) Penaeaceae Melastomataceae Oliniaceae Vochysiaceae s cork origin deep-seated; o r e d endosperm lacking; C resinous, lignans/neolignans, harman alkaloids Zy g o p h y l l a l e s Krameriaceae Zygophyllaceae Eu d i c o t s o s i flw small, G often 3-merous, nectary: intrastaminal disk; Celastraceae Lepidobotryaceae R seeds often with aril (red-orange);

flw pentacyclic, I infl cymose Ce l a s t r a l e s (incl. Hippocrateaceae, Brexiaceae) Parnassiaceae

parts alternating, s Achariaceae Ochnaceae Podostemaceae G connate, d K + C (free), lvs margins toothed; Chrysobalanaceae Hypericaceae Passifloraceae Rhizophoraceae (A polyandrous), flw G often tricarpellate pollen tricolporate; Ma l p i g h i a l e s Clusiaceae Linaceae Salicaceae trihydroxyl-flavonoids lvs often compound, pulvini (sleep movement); Erythroxylaceae Malpighiaceae Picrodendraceae u r o s i flw A5 or multiple, branched style common;

E mucilage cells; oxalates Brunelliaceae Connaraceae Elaeocarpaceae Ox a l i d a l e s Cephalotaceae Cunoniaceae Oxalidaceae 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) Fa b a l e s Fabaceae Polygalaceae Quillajaceae Surianaceae lvs mostly simple with stipules; flw K valvate (and hypanthium) persisting, Cannabaceae Moraceae Ulmaceae carpels with 1 ovule, stigma dry; dihydroflavonols Ro s a l e s Dirachmaceae Rhamnaceae Urticaceae Elaeagnaceae Rosaceae (incl. Cecropiaceae) N fix lvs mostly alternate; flw often unisexual, G mostly inferior, parietal placentation; Anisophyllaceae Coriariaceae Cucurbitaceae cucurbitacines Cucurbitales Begoniaceae Corynocarpaceae Datiscaceae lvs 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 Rhoipteleaceae

II endosperm scanty endosperm

embryo large, large, embryo ectomycorrhiza; tannins, dihydroflavonols Fa g a l e s Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae s

d mostly woody (silica/silicified); lvs alternate, odd-pinnately compound; flw often imperfect, intrastaminal disk; Anacardiaceae Meliaceae Sapindaceae ethereal oils Sa p i n d a l e s Burseraceae Rutaceae Simaroubaceae

u r o s i bark fibrous; hairs often stellate; flw K often valvate,

E Bixaceae Malvaceae (Bombacaceae, Sterculiaceae, Tiliaceae) petals contorted, A often ∞; mucilage, Cistaceae Muntingiaceae Sarcolaenaceae cyclopropenoid fatty acids, flavones a l v a l e s embryo large, M Dipterocarpaceae Neuradaceae Thymelaeaceae endosperm scanty 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 Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae mostly woody; lvs mostly undivided, hydathode teeth; flw often 4-merous, K much smaller than C, persisting, intrastaminal disk, G inferior; fr drupaceous; Cornaceae Hydrangeaceae Loasaceae diverse iridoids Co r n a l e s Grubbiaceae Hydrostachyaceae? Nyssaceae

lvs teeth, theoid; nodes unilacunar; Actinidiaceae Ericaceae Polemoniaceae Sarraceniaceae flw 5-merous, pentacyclic; Balsaminaceae Fouquieriaceae Primulaceae Styracaceae nonhydrolyzable tannins, ellagic acid, hydroquinones Er i c a l e s Clethraceae Lecythidaceae Roridulaceae Theaceae

woody; dioecious; Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae flw small, C valvate, G unilocular; fruit indehiscent; As t e r i d s iridoids (aucubin), gutta Ga r r y a l e s Eucommiaceae Garryaceae (incl. Aucubaceae) mostly sympetalous, nectary gynoecial, late lvs opposite, colleters; sympetaly ovules unitegmic, flw corolla convolute in bud; Apocynaceae (incl. Asclepiadaceae) Loganiaceae endosperm cellular; indole alkaloids; iridoids Ge n t i a n a l e s Gentianaceae Gelsemiaceae Rubiaceae

iridoids common I lvs opposite; nodes 1:1; flw mostly monosymmetric, Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae s A often 2(+2); ethereal oils in gland-headed hairs;

d 6-oxygenated flavones, rosmarinic acid, Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae oligosaccharides: cornoside, verbascoside (acetoside) A=C, La m i a l e s Byblidaceae Martyniaceae Pedaliaceae Stilbaceae epipetalous lvs spiral, simple; nodes unilacunar; polyandry Gesneriaceae Oleaceae Phrymaceae Verbenaceae rare, flw petals plicate; G(2) K persisting; Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae) diverse alkaloids, no iridoids

u a s t e r i o l a n a l e s infl S Hydroleaceae Montiniaceae Sphenocleaceae

cymose E lvs roughly hairy; nodes unilacunar; ellagic acid infl scorpioid; mostly 4 ovules; Boraginaceae Ehretiaceae (incl. Lennoaceae) lacking isokestose, higher inulins, pyrrolizidine alkaloids Bo r a g i n a l e s Cordiaceae Heliotropiaceae Hydrophyllaceae woody; lvs serrate; early flw 1-2 ovules/carpel, C± free, K slightly connate; sympetaly; fr drupe with broad stigma Aquifoliaceae Cardiopteridaceae Stemonuraceae fl small; Aquifoliales HHembryo short woody (except Apiaceae); H lvs often divided; nodes usu. multilacunar; Apiaceae Griseliniaceae Pennantiaceae

Theodor C.H. Cole (Heidelberg) and Hartmut H. Hilger (Berlin) 2008 (Berlin) Hilger H. Hartmut and (Heidelberg) Cole C.H. Theodor infl mostly umbel; drupe or schizocarp p i a l e s

© © (Apiaceae: mericarp/carpophore) A Araliaceae Myodocarpaceae Pittosporaceae

II

s flw often monosymmetric,

______d plunger pollination (long style); Asteraceae Goodeniaceae Pentaphragmataceae - Hypothetical tree based on molecular phylogenetic data (Jan. 2008); branch lengths deliberate, not expressing actual time scale; nodes trilacunar; Calyceraceae Menyanthaceae Rousseaceae position of many characters on tree unclear; if a character is marked as being a potential synapomorphy at a node/for a clade, G inferior inulin, sesquiterpenes, secoiridoids As t e r a l e s this does not mean that all members of that clade possess that character; Campanulaceae (incl. Lobeliaceae) Stylidiaceae - This poster depicts only the largest and most important of the currently accepted approx. 450 families (according to APweb 2008); fr usually 1-few-seeded lvs opposite, often basally connate; nodes 3:3; for family characteristics see: Kubitzki K, ed. (1990 ff.). References: APG II (2003); Judd W et al. (2007); Simpson M (2005); buds with scales; flw often monosymmetric; Adoxaceae Diervillaceae Linnaeaceae Soltis DE et al. (2005); Stevens PF (2008) APweb – www.mobot.org; Watson/Dallwitz (2007) delta-intkey.com/angio/ K persistent in fruit; secoiridoids Valerianaceae

- Thanks to Christoph Dobeš, Ray F. Evert, Peter H. Raven, Douglas E. Soltis, Peter F. Stevens, Maximilian Weigend, Michael Wink u a s t e r i Di p s a c a l e s Caprifoliaceae Dipsacaceae Morinaceae E