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Home , Achariaceae, Actinidiaceae, Alstroemeriaceae, Altingiaceae, Apodanthaceae, Araliaceae

Angiosperm Phylogeny Poster Flowering Systematics

woody; vessels lacking dioecious; flw T5–8, A∞, G5–8, 1/carpel, sac 9-nucleate 1 , 1/1/1 Amborellales Amborellaceae g A herbaceous, aquatic; cambium absent; aerenchyma; flw T4–12, A1–∞, embryo sac 4-nucleate r operculate, perisperm mucilage; (no benzylisoquinolines) 3/6/74 ymphaeales Cabombaceae Hydatellaceae a N

N woody, vessels solitary d flw T>10, A , G ca.9, embryo sac 4-nucleate ∞ Austrobaileyaceae (incl. Illiciaceae) e tiglic acid, aromatic terpenoids 3/5/100 A lvs opposite, interpetiolar nodes swollen

a 1/4/75 e flw small T0–3, A1–5, G1, ovule 1 (apical)/carpel Chloranthales n woody; foliar sclereids a K and C distinct g aromatic terpenoids 2/10/125

r in spherical idioblasts i nodes trilacunar ± herbaceous; lvs two-ranked, base sheathing single adaxial prophyll l (incl. Hydnoraceae) o nodes swollen 4/17/4170

y sesquiterpenes s woody; lvs opposite flw with hypanthium, frequent ethereal oils (pellucid dots) tension +

p anthers often valvate; ovule 1/carpel; embryo large 7/91/2858 aurales Gomortegaceae

L e woody; pith septate; lvs two-ranked with obturator Eupomatiaceae ruminate r features as in 6/128/3140 Degeneriaceae “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 woody infl ± scapose; G often apocarpous Alismataceae Butomaceae Posidoniaceae Scheuchzeriaceae vessels absent often laminar; endosperm helobial Aponogetonaceae Hydrocharitaceae Potamogetonaceae Tofieldiaceae 14/166/4660 eustele Juncaginaceae Ruppiaceae Zosteraceae

sieve tube twining (often); lvs often reticulate

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

steroidal sapogenins/alkaloids 5/21/1050 lvs simple, persistent, entire 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 anthers tetrasporangiate nuclear mostly herbaceous with broad filaments helobial geophytes (bulbs, , ); leaf bases often not sheathing monosulcate pollen monosulcate flw T sometimes spotted, nectaries on T, anthers extrorse sieve tube plastids with seeds ∞ Alstroemeriaceae 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 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 flw pentacyclic Petrosaviales embryo very small P 3-merous, A opp. P often geophytes (incl. Agapanthaceae, Alliaceae) Hypoxidaceae Iridaceae filaments narrow fr or anthers broadly attached coat obliterated or with phytomelan (incl. Agavaceae, Hyacinthaceae, Ruscaceae) Lanariaceae Orchidaceae 14/1122/36205 septal nectary woody, often monopodial Tecophilaeaceae (incl. Xanthorrhoeaceae, Hemerocallidaceae) single 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 stem-borne numerous alkaloids herbs (mostly); lvs grassy; siliceous – (in most) Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae

flw often anemophilous, minute, chaffy, without nectaries 15/997/18875 Rapateaceae Typhaceae (incl. Sparganiaceae) infl thyrsus of scorpioid cymes flw monosymmetric or not, few fertile Commelinaceae Haemodoraceae Hanguanaceae phenylphenalenones 5/68/812 Philydraceae Pontederiaceae UV-fluorescing cell walls herbs, rhizomatous, large-leafed; pseudostem common (ferulic/coumaric acids) flw irregular/monosymmetric, septal nectaries silicic acid in A often strongly modified/reduced, G inferior; seeds often arillate Cannaceae Heliconiaceae cuticular waxes often in rodlets silicic acid 8/92/~2500 ingiberales aggregated into scallops Z Lowiaceae aquatic; herbaceous; monoecious lvs whorled, no pellucid dots; vessels lacking flw T0 or 9–10*, A1, G1, 1 apical ovule/carpel 1/1/6 pollen inaperturate, pollen tube branched, hydrophilous Ceratophyllales lvs often divided flw parts whorled, P single or multiple whorls Berberidaceae Eupteleaceae G apocarpous/paracarpous, superior ethereal berberines 7/199/4510 Circaeasteraceae Lardizabalaceae 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 woody; vessels lacking flw T 0, A∞, G>5 laterally connate with abaxial nectaries 1/2/2 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 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 ; fr usu. follicles 1/10/300 s lvs with glandular teeth; often hypanthium, apically unfused carpels, decurrent Cynomoriaceae fr mostly dry, dehiscent pollen tricolpate myricetin, flavonols Cercidiphyllaceae Daphniphyllaceae Paeoniaceae protandry common 15/112/2500 flw K/C/P opp A Grossulariaceae filaments rather narrow often tendrillar vines; lvs often divided and with glandular teeth nodes 3:3 A epipetalous, ovules 2/carpel; raphides, pearl glands benzylisoquinolines berries stomata anomocytic 1/14/850 itales microsporogenesis V simultaneous cork origin deep-seated endosperm lacking resinous, /neolignans, harman alkaloids Krameriaceae 2/24/345 lvs often compound, pulvini (sleep movement) stipules flw A5 or multiple, branched style common Brunelliaceae Connaraceae Elaeocarpaceae mucilage cells Oxalidaceae oxalates 7/60/1845 Cephalotaceae gallic acids gallic S

and infl cymose, flw small G often 3-merous, nectary often intrastaminal disk u COM seeds often arillate (red-orange) or winged (incl. Hippocrateaceae, Brexiaceae, ) p 2/94/1355 ellagic ellagic e habits and extremely diverse Achariaceae Euphorbiaceae Podostemaceae r F lvs margins toothed; pedicels often articulated Hypericaceae Passifloraceae Rhizophoraceae flw G3 (often) c E r a 36/716/16065 Clusiaceae Linaceae Phyllanthaceae u o b flwoften “papilionaceous”: wing, standard, keel, C clawed, G1 (most), A10 (most) Erythroxylaceae Malpighiaceae Picrodendraceae o fr pod d s R i with nodule bacteria r 4/754/20140 i i o d diverse alkaloids, NP amino acids, lectins (in ) Fabaceae Quillajaceae Surianaceae e s lvs mostly simple with stipules c d s flw K valvate (and hypanthium) persisting Barbeyaceae o s i stigma dry, ovule 1/carpel Cannabaceae d N fix dihydroflavonols 9/261/7725 osales t R Dirachmaceae (incl. Cecropiaceae) s embryo large lvs mostly alternate s endosperm scanty flw often unisexual, G mostly inferior parietal placentation Apodanthaceae Begoniaceae Corynocarpaceae Datiscaceae 7/109/2935 Anisophyllaceae Coriariaceae Tetramelaceae mostly ; lvs mostly undivided; flw small, unisexual anemophilous, thus T reduced or lacking, G mostly inferior Fagaceae infl spikes or ; fr mostly nuts; seed 1 Ticodendraceae ectomycorrhiza; , dihydroflavonols 7/33/1005 Casuarinaceae Juglandaceae Nothofagaceae stems jointed at nodes (in some); lvs with glandular teeth flw A obdiplostemonous, nectary outside A; fr capsule (incl. Ledocarpaceae, , ) ethereal oils, ellagic acid 2(5)/17/897 lvs opposite, colleters (glandular hair on adaxial surface of base) Combretaceae Penaeaceae (incl. Oliniaceae) stipules small (if any), cork deep seated flw 5-merous flw K valvate, persisting; A incurved in , ovary inferior, ovules many Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae)

parts whorled endosperm scanty, scaly ; flavonols, myricetin 9/380/13005 yrtales K + C free P M Melastomataceae (incl. Memecylaceae) Onagraceae diplostemonous e woody pollen tricolporate stylodia free Crossosomataceae Stachyuraceae G connate n hypanthium with nectary 7/12/66 rossosomatales 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, 1/3/49 Picramniales 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 Biebersteiniaceae Meliaceae Rutaceae 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 endo- sperm bark fibrous; hairs often stellate Bixaceae (incl. Bombacaceae, , ) scanty Liverworts flw K often valvate, contorted, A often∞ ; mucilage Cistaceae Cytinaceae Muntingiaceae Sarcolaenaceae cyclopropenoid fatty acids, flavones 10/338/6005 Dipterocarpaceae Neuradaceae Sphaerosepalaceae Thymelaeaceae woody or herbs infl racemose; flw often 4-merous, petals often clawed Bataceae Caricaceae Limnanthaceae Salvadoraceae myrosin cells, 18/405/5035 rassicales Moringaceae Tovariaceae B Koeberliniaceae Tropaeolaceae woody; (semi-)parasites; mycorrhiza absent; lvs margins entire flw A epipetalous; often simple, valvate, persisting Tracheophytes (incl. horsetails) carpels/ovaries/ovules often reduced, placentation free-central; fr , one-seeded Balanophoraceae Misodendraceae Opiliaceae Schoepfiaceae polyacetylenes, triterpene sapogenins, silicic acid 13/151/1992 antalales Ginkgo S Loranthaceae Viscaceae woody Ephedra Welwitschia lvs stomata cyclocytic, petiole bundles annular

Gnetum fr indehiscent (more or less fleshy) 2/3/4 Seed calcium oxalate as crystals erberidopsidales Aextoxicaceae Berberidopsidaceae 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 betalains or (latter, e.g., in Caryophyllaceae) 37/749/11620 Monocots S 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 Malvids e 6/51/590 diverse iridoids ornales r C Curtisiaceae Hydrostachyaceae amiids seed L lvs teeth often theoid; nodes unilacunar a exo- testal flw 5-merous, pentacyclic Fouquieriaceae Campanulids s nonhydrolyzable tannins, ellagic acid embryo triterpenoids, hydroquinones 22/346/11545 t long Roridulaceae e trop. trees; lvs exstipulate/entire; nodes often unilacunar Myrsinaceae Theophrastaceae pedicels articulated; style short; embryo long r endosperm copious i indole and quinoline alkaloids (camptothecin) 2/24/202 Icacinales Oncothecaceae d trop. trees; few shared features; nodes usu. trilacunar alkaloids, iridoids Theodor C. H. Cole, Dipl. Biol. s : lvs alternate; flw bisexual, fragrant; anthers sagittate; pollen sacs moniliform, dehisce longitudinally

1/11/55 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 Institute of Biology – Botany fr indehiscent Eucommiaceae (incl. Aucubaceae) iridoids (aucubin), gutta 2/3/18 arryales Freie Universität Berlin e L G Altensteinstr. 6, D-14195 Berlin, r a lvs opposite; nodes 1:1 flw mostly monosymmetric i m A often 2(+2); gland-headed hairs with radially arranged cells d ii 6-oxygenated flavones, oligosaccharides cornoside, (acetoside) 24/1059/23810 Byblidaceae s d herbaceous; lvs opposite, glandular-pubescent Peter F. Stevens, Ph.D. flw < 5 mm across, K valvate, G(3) inf, nectary s fr septicidal capsule, K/C/A persistent; seeds minute mostly sympetalous Missouri Botanical Garden (MoBot) to ; iridoids 1/1/8 C enclosing Vahliales Vahliaceae St. Louis, MO 63166-0299, USA A and G in bud lvs spiral, simple; nodes unilacunar and G(2) flw petals plicate style single, long C K persistent in forming (incl. Cuscutaceae) (incl. Nolanaceae) University of Missouri – St. Louis nectary gynoecial A=C alkaloids diverse, no iridoids 5/165/4125 ovules unitegmic polyandry distinct Hydroleaceae Sphenocleaceae Department of Biology rare tube St. Louis, MO 63121- 4499, USA endosperm cellular lvs opposite iridoids + infl A interpetiolar stipules or nodal line/ridge, colleters cymose epipetalous flw corolla convolute in bud (incl. Asclepiadaceae) fr drupe indole alkaloids, iridoids 5/1121/19915 ellagic acid lacking lvs roughly hairy; nodes unilacunar infl scorpioid mostly 4 ovules Codonaceae Coldeniaceae Cordiaceae Ehretiaceae (+ Lennoaceae)

COLE TCH, HILGER HH, STEVENS PF (2019) Angiosperm Phylogeny Poster – Systematics isokestose, higher inulins, pyrrolizidine alkaloids 6-11/150/3095 Heliotropiaceae Hydrophyllaceae Namaceae Wellstediaceae • hypothetical based on molecular phylogenetic (chiefly ) data (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 • 64 orders and around 420 families currently recognized by APG IV (some minor families omitted here) Aquifoliaceae fr usu. drupe with broad stigma 5/21/536 quifoliales • contrary to APG, but in compliance with APweb and other seminal sources, we here recognize several families within Boraginales A • branch lengths deliberate, not expressing actual time scale herbs or ; nodes trilacunar early • the characters listed do not necessarily apply to all members of a clade flw often monosymmetric Pentaphragmataceae sympetaly plunger sec. device (long style) • position of various characters on the tree uncertain flw small inulin, sesquiterpenes, secoiridoids 11/1743/26870 sterales for characteristics see: FGVP, Kubitzki K, ed. (1990 ff) embryo A • (incl. Lobeliaceae) • orders (and higher ranks) are linked to the according pages on APweb short mostly woody • numbers set in gray next to orders refer to families, genera, and species infl racemose, C free

anthers basifixed, nectary disk 1/9/130 * : the alleged could be (issue being addressed in ongoing and debate) Escalloniales This poster is now available in 30 languages (please refer to the authors’ pages on ResearchGate) C woody; evergreen References: Stevens PF (2020) APweb – www.mobot.org/MOBOT/research/APweb a nodes 1:1 2/14/79 (incl. ) APG IV (2016); Judd W et al. (2016); Li HT et al. (2019); Simpson M (2020); Soltis DE et al. (2014); Watson/Dallwitz (2018) m flw polysymmetric, anthers basifixed ) Acknowledgements to our contributors, translators, and consultants: Rubina Abid, Julien B. Bachelier, Przemysław Baranow, Zoltán Barina, Fernanda Antunes Carvalho, p woody (except ) Christoph Dobeš, Johannes Enroth, Ray F. Evert, Mohamed Fennane, Marc Gottschling, Zigmantas Gudžinskas, Michal Hroneš, Akitoshi Iwamoto, Chen-Kun Jiang, Anna Kagiampaki, lvs often divided; nodes usu. multilacunar G Kent Kainulainen, Elizabeth A. Kellogg, Sangtae Kim, Aslı Doğru-Koca, Nikos Krigas, Sanjay Kumar, Ľuboš Majeský, Pavol Mártonfi, Diego Medan, Sergei L. Mosyakin, a infl mostly ; fr drupe or schizocarp Apiaceae Griseliniaceae Pennantiaceae CC-BY

( Sofi Mursidawati, Mayur Nandikar, Dashzeveg Nyambayar, Anastasiya V. Odintsova, Richard G. Olmstead, Batlai Oyuntsetseg, Peter H. Raven, Yasaman Salmaki, Federico Selvi, inferior (Apiaceae: mericarp/carpophore) 7/494/5489 Paramjit Singh, Douglas E. Soltis, Bach The Tran, Pramote Triboun, Zoya M. Tsymbalyuk, Orzimat T. Turginov, Magsar Urgamal, Maximilian Weigend, Michael Wink, Shahin Zarre piales n fr A u usually woody 1–few-seeded infl racemose, flw 4-merous, filaments stout l capsule septicidal 1/3/36 Angiosperm Tracheophyte Paracryphiales i lvs opposite, often basally connate Phylogeny Phylogeny Phylogeny d with scales; flw often monosymmetric K persistent in fruit Poster Poster Poster s secoiridoids 2/46/1090 (incl. ) © The Authors, 2019/3 © The Diervillaceae Linnaeaceae Morinaceae Valerianaceae