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 1/1/1 Amborellales Amborellaceae g 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) 3/6/74 ymphaeales Cabombaceae Hydatellaceae Nymphaeaceae a N N woody, vessels solitary d flw T>10, A , G ca.9, embryo sac 4-nucleate ∞ Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae e tiglic acid, aromatic terpenoids 3/5/100 Austrobaileyales A lvs opposite, interpetiolar stipules flw small T0–3, A1–5, G1, 1 apical ovule/carpel

a 1/4/75 Chloranthaceae e nodes swollen Chloranthales n woody; foliar sclereids a K and C distinct g aromatic terpenoids 2/10/125 Canellales Canellaceae Winteraceae r idioblasts spherical in i nodes trilacunar ± herbaceous; lvs two-ranked, leaf base sheathing single adaxial prophyll l Aristolochiaceae (incl. Hydnoraceae) Piperaceae Saururaceae o nodes swollen 4/17/4170 Piperales

y sesquiterpenes s woody; lvs opposite

flw with hypanthium, staminodes frequent Calycanthaceae Hernandiaceae Monimiaceae

tension wood + wood tension

(pellucid dots) (pellucid ethereal oils ethereal

p anthers often valvate; carpels with 1 ovule; embryo large 7/91/2858 aurales Gomortegaceae Lauraceae Siparunaceae

L e Magnoliids woody; pith septate; lvs two-ranked ovules with obturator Annonaceae Eupomatiaceae Magnoliaceae endosperm ruminate r features as in 6/128/3140 Magnoliales Degeneriaceae Himantandraceae Myristicaceae “Early Angiosperms” infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules m ovules atropous, seeds with epidermal perisperm and copious endosperm s ethereal oils 1/1/2-4 Acorales Acoraceae mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules woody infl ± scapose; flw G apocarpous; placentation often laminar Alismataceae Butomaceae Posidoniaceae Scheuchzeriaceae vessels absent endosperm helobial; embryo large/green Aponogetonaceae Hydrocharitaceae Potamogetonaceae Tofieldiaceae 14/166/4660 Alismatales eustele Araceae Juncaginaceae Ruppiaceae Zosteraceae

sieve tube plastids twining vines (often); lvs often reticulate

endosperm triploid endosperm embryo sac 8-nucleate sac embryo with starch grains alkaloids benzylisoquinoline ovary often inferior, style short, branched

steroidal sapogenins/alkaloids 5/21/1050 lvs simple, persistent, entire Dioscoreales Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae Thismiaceae flw strobilar, perfect, parts free Monocots Ca oxalate P parts varying, often in threes, raphides some woody (with terminally tufted lvs) scattered bundles in stem

weakly differentiated inflsometimes with spathe + spadix 5/36/1345 no secondary thickening endosperm Cyclanthaceae Pandanaceae Triuridaceae Velloziaceae anthers tetrasporangiate nuclear Pandanales mostly herbaceous stamen with broad filaments helobial often geophytes (bulbs, tubers, rhizomes); leaf bases often not sheathing pollen monosulcate pollen monosulcate flw T sometimes spotted, nectaries on T, anthers extrorse sieve tube plastids with seeds many Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae G apocarpous (style short in most) protein crystals phytomelan lacking; fructans in stems, chelidonic acid 10/67/1558 iliales compitum (if present) extragynoecial L Colchicaceae Liliaceae Petermanniaceae Smilacaceae sympodial branching nectaries absent stem with ring of bundles lvs parallel-veined, entire siphonogamy fr follicle

no glandular teeth 1/2/3 double fertilization > endosperm East Asia Petrosaviaceae flw pentacyclic Petrosaviales embryo very small P 3-merous, A opp. P often geophytes Amaryllidaceae (incl. Agapanthaceae, Alliaceae) Hypoxidaceae Iridaceae filaments narrow fr capsule or berry anthers broadly attached seed coat obliterated or with phytomelan Asparagaceae (incl. Agavaceae, Hyacinthaceae, Ruscaceae) Lanariaceae Orchidaceae 14/1122/36205 Asparagales septal nectary woody, often monopodial Tecophilaeaceae Asphodelaceae (incl. Xanthorrhoeaceae, Hemerocallidaceae) single cotyledon lvs often palmately or pinnately pseudocompound, reduplicate-plicate extrafloral nectaries extrafloral radicle not persistent intense primary growth, large apical meristem, infl often with spathe

1/188/2585 Arecaceae Dasypogonaceae stem-borne roots numerous alkaloids Arecales mostly herbaceous; lvs grassy; epidermis siliceous mycorrhiza – (in most) Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae

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; seeds often arillate Cannaceae Heliconiaceae Marantaceae Strelitziaceae cuticular waxes often in rodlets silicic acid 8/92/~2500 ingiberales aggregated into scallops Z 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 1/1/6 Ceratophyllaceae pollen inaperturate, pollen tube branched, hydrophilous Ceratophyllales lvs often divided flw parts whorled, P single or multiple whorls Berberidaceae Eupteleaceae Menispermaceae G apocarpous/paracarpous, superior Ranunculaceae ethereal berberines 7/199/4510 Ranunculales Circaeasteraceae Lardizabalaceae Papaveraceae oils mostly woody not in flw T often 4-merous Nelumbonaceae Platanaceae Proteaceae Sabiaceae idioblasts A epitepalous, connectives sometimes with apical appendage 4/85/1750 Proteales woody; vessels lacking flw T missing, A∞, G>5 laterally connate with abaxial nectaries 1/2/2 Trochodendraceae recepta- fr aggregate of follicles Trochodendrales cular E nectary mostly woody; lvs evergreen, stomata cyclocytic mostly monoecious, flw unisexual, T ± uniform or missing

common 1/7/120 Buxaceae (incl. Haptanthaceae) u pregnane pseudoalkaloids Buxales d lvs toothed, sec. veins palmate i dioecious, flw unisexual, T small to lacking 2/2/50 unnerales Gunneraceae Myrothamnaceae c ellagic acid G o absent mostly woody; lvs if veins strong, proceed to apex of teeth flw mostly K5, persisting, mostly A∞, G mostly slightly connate Dilleniaceae t seeds often with aril; fr usually follicles 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 Crassulaceae Grossulariaceae Iteaceae Saxifragaceae filaments rather narrow often tendrillar vines; lvs often divided and with glandular teeth nodes 3:3 A epipetalous, 2 ovules per carpel; raphides, pearl glands benzylisoquinolines berries stomata anomocytic 1/14/850 itales Vitaceae microsporogenesis V simultaneous cork origin deep-seated endosperm lacking resinous, lignans/neolignans, harman alkaloids Krameriaceae Zygophyllaceae 2/24/345 Zygophyllales lvs often compound, pulvini (sleep movement) stipules flw A5 or multiple, branched style common Brunelliaceae Connaraceae Elaeocarpaceae mucilage cells Oxalidaceae oxalates 7/60/1845 Oxalidales Cephalotaceae Cunoniaceae Huaceae gallic acids gallic S

and infl cymose, flw small G often 3-merous, nectary often intrastaminal disk u COM clade seeds often arillate (red-orange) or winged Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae) Lepidobotryaceae p 2/94/1355 Celastrales ellagic ellagic e habits and habitats extremely diverse Achariaceae Euphorbiaceae Rafflesiaceae Ochnaceae Podostemaceae r F lvs margins toothed Chrysobalanaceae Hypericaceae Passifloraceae Rhizophoraceae flw G often tricarpellate c E r a 36/716/16065 Malpighiales Clusiaceae Linaceae Phyllanthaceae Salicaceae u o b flwoften “papilionaceous”: wing, standard, keel, C clawed, G1 (most), A10 (most) Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae o fr pod d s R i symbiosis with root nodule bacteria r 4/754/20140 i i o d diverse alkaloids, NP amino acids, lectins (in Fabaceae) Fabales Fabaceae Polygalaceae Quillajaceae Surianaceae e s lvs mostly simple with stipules c d s flw K valvate (and hypanthium) persisting Barbeyaceae Elaeagnaceae Rosaceae o s i carpels with 1 ovule, stigma dry Cannabaceae Moraceae Ulmaceae d N fix dihydroflavonols 9/261/7725 osales t R Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae) s embryo large lvs mostly alternate s endosperm scanty flw often unisexual, G mostly inferior parietal placentation Apodanthaceae Begoniaceae Corynocarpaceae Datiscaceae 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 Betulaceae Fagaceae Myricaceae infl spikes or catkins; fr 1-seeded, mostly nuts Ticodendraceae ectomycorrhiza; tannins, dihydroflavonols 7/33/1005 Fagales Casuarinaceae Juglandaceae Nothofagaceae stems sometimes jointed at nodes; lvs with glandular teeth flw A obdiplostemonous, nectary outside A; fr capsule Geraniaceae Francoaceae (incl. Ledocarpaceae, Melianthaceae, Vivianiaceae) ethereal oils, ellagic acid 2(5)/17/897 Geraniales lvs opposite, colleters (glandular hair on adaxial surface of petiole base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae) stipules small (if any), cork deep seated flw 5-merous flw K valvate, persisting, A incurved in bud, ovary inferior, ovules many Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae)

parts whorled endosperm scanty, scaly bark, flavonols, myricetin 9/380/13005 yrtales K + C free P M Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae diplostemonous e woody pollen tricolporate stylodia free Crossosomataceae Stachyuraceae G connate n hypanthium with nectary 7/12/66 rossosomatales Strasburgeriaceae style + t C Geissolomataceae Staphyleaceae endosperm nuclear trees, dioecious, lvs compound a M extrafloral nectaries, staminate flw: A = and opposite C p a bark bitter, anthraquinones 1/3/49 Picramniales Picramniaceae e l mostly woody (silica/silicified) lvs often alternate, odd-pinnately compound t v flw often imperfect, intrastaminal disk Anacardiaceae Burseraceae Kirkiaceae Nitrariaceae Sapindaceae a i ethereal oils 9/479/6550 Sapindales Biebersteiniaceae Meliaceae Rutaceae Simaroubaceae l d vessel elements with scalariform perforations; mucilage cells a s embryo lvs margins toothed, stipules cauline

4/6/24 Dipentodontaceae Gerrardinaceae Petenaeaceae Tapisciaceae e large flw small, A = and opposite K, ovules 1-2/carpel Huerteales endo- sperm bark fibrous; hairs often stellate Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae) scanty Liverworts flw K often valvate, petals contorted, A often∞ ; mucilage Cistaceae Cytinaceae Muntingiaceae Sarcolaenaceae cyclopropenoid fatty acids, flavones 10/338/6005 Malvales Mosses Dipterocarpaceae Neuradaceae Sphaerosepalaceae Thymelaeaceae woody or herbs Hornworts flw often 4-merous, often clawed petals, infl racemose Bataceae Caricaceae Limnanthaceae Salvadoraceae myrosin cells, glucosinolates 18/405/5035 rassicales Brassicaceae Cleomaceae Moringaceae Tovariaceae Lycophytes B Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae woody; (semi-)parasites; mycorrhiza absent; lvs margins entire Ferns flw A epipetalous; perianth often simple, valvate, persisting Tracheophytes (incl. horsetails) carpels/ovaries/ovules often reduced, placentation free-central; fr drupe, one-seeded Balanophoraceae Misodendraceae Opiliaceae Schoepfiaceae Cycads polyacetylenes, triterpene sapogenins, silicic acid 13/151/1992 antalales Ginkgo S Loranthaceae Olacaceae Santalaceae Viscaceae woody Ephedra Welwitschia lvs stomata cyclocytic, petiole bundles annular

Gnetum fr indehiscent (more or less fleshy) 2/3/4 Gymnosperms Seed Conifers calcium oxalate as crystals erberidopsidales Aextoxicaceae Berberidopsidaceae Plants B ANA grade mostly herbaceous; without mycorrhiza Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae G often unilocular with free-central placentation Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae Angiosperms Magnoliids pollen colpate, surface spiny Chenopodiaceae Droseraceae Nyctaginaceae Simmondsiaceae betalains or anthocyanins (latter, e.g., in Caryophyllaceae) 37/749/11620 Monocots S Caryophyllales Basellaceae Drosophyllaceae Phytolaccaceae Talinaceae u Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae Fabids mostly woody; lvs mostly undivided, hydathode teeth p flw often 4-merous, K much smaller than C, persisting intrastaminal disk, G inferior; fr drupaceous Cornaceae Grubbiaceae Loasaceae Malvids e 6/51/590 Rosids diverse iridoids ornales r C Curtisiaceae Hydrangeaceae Hydrostachyaceae Nyssaceae amiids seed L lvs teeth often theoid; nodes unilacunar Asterids a exo- Actinidiaceae Ericaceae Pentaphylacaceae Sarraceniaceae testal flw 5-merous, pentacyclic Balsaminaceae Fouquieriaceae Polemoniaceae Styracaceae Campanulids s nonhydrolyzable tannins, ellagic acid embryo triterpenoids, hydroquinones 22/346/11545 Clethraceae Lecythidaceae Primulaceae Symplocaceae t long Ericales Diapensiaceae Marcgraviaceae Roridulaceae Theaceae e trop. trees; lvs exstipulate/entire; nodes often unilacunar Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae pedicels articulated; style short; embryo long r endosperm copious i indole and quinoline alkaloids (camptothecin) 2/24/202 Icacinales Oncothecaceae Icacinaceae d trop. trees; few shared features; nodes usu. trilacunar alkaloids, iridoids Theodor C. H. Cole, Dipl. Biol. s Metteniusa: lvs alternate; flw bisexual, fragrant; anthers sagittate; pollen sacs moniliform, dehisce longitudinally

1/11/55 Metteniusaceae A late infl axillary cyme;G (5) unilocular; NE Andes, cloud montane forest Metteniusales Hartmut H. Hilger, Prof. Dr. sympetaly s woody; dioecious Dahlem Centre of Plant Sciences (DCPS) t flw small, C valvate, G unilocular fr indehiscent Eucommiaceae Garryaceae (incl. Aucubaceae) Institute of Biology – Botany e iridoids (aucubin), gutta 2/3/18 Garryales Freie Universität Berlin L r a lvs opposite; nodes 1:1 Altensteinstr. 6, D-14195 Berlin, Germany flw mostly monosymmetric Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae i m A often 2(+2); gland-headed hairs with radially arranged cells Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae d ii 6-oxygenated flavones, oligosaccharides cornoside, verbascoside (acetoside) 24/1059/23810 Lamiales Byblidaceae Martyniaceae Pedaliaceae Stilbaceae s d herbaceous; lvs opposite, glandular-pubescent Calceolariaceae Gesneriaceae Oleaceae Phrymaceae Verbenaceae flw < 5 mm across, K valvate, G(3) inf, nectary s fr septicidal capsule, K/C/A persistent; seeds minute Peter F. Stevens, Ph.D. mostly sympetalous Africa to India; iridoids 1/1/8 ahliales Vahliaceae Missouri Botanical Garden (MoBot) C enclosing V A and G in bud lvs spiral, simple; nodes unilacunar St. Louis, MO 63166-0299, USA G(2) flw petals plicate style single, long C K persistent in fruit and forming Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae) nectary gynoecial A=C alkaloids diverse, no iridoids 5/165/4125 ovules unitegmic polyandry distinct olanales University of Missouri – St. Louis tube S Hydroleaceae Montiniaceae Sphenocleaceae endosperm cellular rare lvs opposite Department of Biology iridoids + infl A interpetiolar stipules or nodal line/ridge, colleters St. Louis, MO 63121 - 4499, USA cymose epipetalous flw corolla convolute in bud Apocynaceae (incl. Asclepiadaceae) Loganiaceae fr drupe indole alkaloids, iridoids 5/1121/19915 entianales seed single G Gentianaceae Gelsemiaceae Rubiaceae ellagic acid lvs roughly hairy; nodes unilacunar lacking infl scorpioid mostly 4 ovules Boraginaceae Codonaceae Coldeniaceae Cordiaceae Ehretiaceae (+ Lennoaceae)

isokestose, higher inulins, pyrrolizidine alkaloids 6-11/150/3095 Boraginales Heliotropiaceae Hydrophyllaceae Namaceae Wellstediaceae COLE TCH, HILGER HH, STEVENS PF (2019) Angiosperm Phylogeny Poster – Flowering Plant Systematics • hypothetical tree based on molecular phylogenetic (chiefly plastid) data (May 2019) woody; lvs serrate • phylogeny, classification, and features chiefly followAPweb , APG, and PPA (Plastid Phylogenomic Angiosperm see Li HT et al. 2019) flw 1-2 ovules/carpel, C ± free, K slightly connate Aquifoliaceae Cardiopteridaceae Stemonuraceae • 64 orders and around 420 families currently recognized by APG IV (some minor families omitted here) fr usu. drupe with broad stigma 5/21/536 Aquifoliales • contrary to APG, but in compliance with APweb and other seminal sources, we here recognize several families within Boraginales herbs or shrubs; nodes trilacunar • branch lengths deliberate, not expressing actual time scale early flw often monosymmetric Asteraceae Goodeniaceae Pentaphragmataceae • the characters listed do not necessarily apply to all members of a clade sympetaly plunger sec. pollination device (long style) flw small Calyceraceae Menyanthaceae Rousseaceae inulin, sesquiterpenes, secoiridoids 11/1743/26870 • position of various characters on the tree uncertain embryo Asterales • for family characteristics see: FGVP, Kubitzki K, ed. (1990 ff) short (incl. Lobeliaceae) mostly woody Campanulaceae Stylidiaceae • orders (and higher ranks) are linked to the according pages on APweb infl racemose, C free numbers set in gray next to orders refer to families, genera, and species • anthers basifixed, nectary disk 1/9/130 scalloniales Escalloniaceae * Ceratophyllum: the alleged tepals could be bracts (issue being addressed in ongoing research and debate) C E This poster is now available in 25 languages (please refer to the authors’ pages on ResearchGate) woody; evergreen a nodes 1:1 References: Stevens PF (2019) APweb – www.mobot.org/MOBOT/research/APweb flw polysymmetric, anthers basifixed 2/14/79 Bruniaceae Columelliaceae (incl. Desfontainia) APG IV (2016); Judd W et al. (2016); Li HT et al. (2019); Simpson M (2010); Soltis DE et al. (2014); Watson/Dallwitz (2018) m Bruniales ) p woody (except Apiaceae) Acknowledgements to our contributors, translators, and consultants: Rubina Abid, Julien B. Bachelier, Przemysław Baranow, Zoltán Barina, Fernanda Antunes Carvalho, Christoph Dobeš, lvs often divided; nodes usu. multilacunar Ray F. Evert, Mohamed Fennane, Marc Gottschling, Zigmantas Gudžinskas, Akitoshi Iwamoto, Chen-Kun Jiang, Anna Kagiampaki, Kent Kainulainen, Elizabeth A. Kellogg, Sangtae Kim, G Apiaceae Griseliniaceae Pennantiaceae CC-BY a infl mostly umbel; fr drupe or schizocarp ( inferior Aslı Doğru-Koca, Nikos Krigas, Sanjay Kumar, Diego Medan, Sergei L. Mosyakin, Sofi Mursidawati,Dashzeveg Nyambayar, Anastasiya V. Odintsova, Richard G. Olmstead, Batlai Oyuntsetseg, (Apiaceae: mericarp/carpophore) 7/494/5489 piales Araliaceae Myodocarpaceae Pittosporaceae Peter H. Raven, Yasaman Salmaki, Federico Selvi, Paramjit Singh, Douglas E. Soltis, Bach The Tran, Pramote Triboun, Zoya M. Tsymbalyuk, Magsar Urgamal, Maximilian Weigend, Michael Wink, Shahin Zarre n fr A u usually woody 1–few-seeded infl racemose, flw 4-merous, filaments stout l capsule septicidal 1/3/36 Paracryphiaceae Angiosperm Tracheophyte Bryophyte Paracryphiales i lvs opposite, often basally connate Phylogeny Phylogeny Phylogeny d buds with scales; flw often monosymmetric Adoxaceae K persistent in fruit Poster Poster Poster s 2/46/1090 © The Authors, 2019/3 © The secoiridoids Dipsacales Caprifoliaceae (incl. Dipsacaceae Diervillaceae Linnaeaceae Morinaceae Valerianaceae)

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