Angiosperm Phylogeny Flowering Systematics

woody, vessels lacking; dioecious; flw T5–8, A∞, G5–8, 1 /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 ∞ 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, base sheathing o single adaxial prophyll; swollen nodes Aristolochiaceae (incl. Hydnoraceae) Piperaceae Saururaceae 4/17/4170 Piperales

y sesquiterpenes

s woody; lvs opposite; flw with hypanthium, 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; 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; 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 often inferior, style short, branched; steroidal sapogenins/alkaloids ioscoreales Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae Thismiaceae

helobial 5/21/1050 D with broad filaments mostly herbaceous pollen monosulcate pollen monosulcate some woody (with terminally tufted ) 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 often geophytes P 3-merous, A opp. P incl. Agapanthaceae, Alliaceae filaments narrow or berry Amaryllidaceae ( ) Hypoxidaceae Iridaceae anthers broadly attached 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 Arecales Arecaceae Dasypogonaceae mostly herbaceous; 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 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, decurrent Altingiaceae Cynomoriaceae Peridiscaceae fr mostly dry, dehiscent pollen tricolpate myricetin, flavonols Cercidiphyllaceae Daphniphyllaceae Paeoniaceae protandry common 15/112/2500 flw K/C/P opp A often tendrillar vines; lvs often divided and with glandular teeth Grossulariaceae Iteaceae 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 (incl. Hippocrateaceae, Brexiaceae, ) Lepidobotryaceae 2/94/1355 gallic acids gallic S and lvs often compound, pulvini (sleep movement) u COM flw A5 or multiple, branched style common Brunelliaceae Connaraceae Elaeocarpaceae p mucilage cells; oxalates xalidales Oxalidaceae 7/60/1845 ellagic ellagic O Cephalotaceae Cunoniaceae e r habits and habitats extremely diverse Achariaceae Euphorbiaceae Rafflesiaceae Podostemaceae F lvs margins toothed Chrysobalanaceae Hypericaceae 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 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 ; 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 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 flw 5-merous lvs opposite, colleters (glandular hair on adaxial surface of base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae) parts whorled stipules small (if any), cork deep seated (incl. Punicaceae, Sonneratiaceae, Trapaceae) K + C free P flw K valvate, persisting, A incurved in , ovary inferior, ovules many Lythraceae diplostemonous endosperm scanty, scaly bark, flavonols, myricetin yrtales e 9/380/13005 M Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae pollen tricolporate G connate n woody style + stylodia free Crossosomataceae Stachyuraceae t hypanthium with nectary Strasburgeriaceae endosperm nuclear 7/12/66 Crossosomatales Geissolomataceae Staphyleaceae a trees, dioecious, lvs compound, extrafloral nectaries p staminate flw: A = and opposite C M Picramniaceae

1/3/49 icramniales e a bark bitter, anthraquinones P t l mostly woody (silica/silicified) Anacardiaceae Burseraceae Kirkiaceae Nitrariaceae Sapindaceae a lvs often alternate, odd-pinnately compound v flw often imperfect, intrastaminal disk,ethereal oils 9/479/6550 Sapindales Biebersteiniaceae Meliaceae Rutaceae Simaroubaceae l i vessel elements with scalariform perforations; mucilage cells lvs margins toothed, stipules cauline a embryo Dipentodontaceae Gerrardinaceae Petenaeaceae Tapisciaceae d flw small, A = and opposite K, ovules 1-2/carpel uerteales e large 4/6/24 H s (incl. Bombacaceae, Sterculiaceae, Tiliaceae) endo- bark fibrous; hairs often stellate Bixaceae Malvaceae sperm scanty flw K often valvate, contorted, A often∞ ; mucilage Cistaceae Cytinaceae Muntingiaceae Sarcolaenaceae Liverworts cyclopropenoid fatty acids, flavones alvales 10/338/6005 M Dipterocarpaceae Neuradaceae Sphaerosepalaceae Thymelaeaceae Mosses woody or herbs; flw often 4-merous Bataceae Caricaceae Limnanthaceae Salvadoraceae Hornworts often clawed petals, infl racemose myrosin cells, glucosinolates Brassicaceae Cleomaceae Moringaceae Tovariaceae 18/405/5035 Brassicales Lycophytes Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae woody; lvs stomata cyclocytic

Ferns petiole bundles annular; fr indehiscent, more or less fleshy Tracheophytes (incl. horsetails) calcium oxalate as crystals Aextoxicaceae Berberidopsidaceae 2/3/4 erberidopsidales Cycads B Ginkgo woody; (semi-)parasites; mycorrhiza absent; lvs margins entire flw A epipetalous; often simple, valvate, persisting Ephedra Balanophoraceae Misodendraceae Opiliaceae Schoepfiaceae Welwitschia carpels/ovaries/ovules often reduced, placentation free-central; fr , one-seeded

Gnetum polyacetylenes, triterpene sapogenins, silicic acid 13/151/1992 antalales Loranthaceae Olacaceae Santalaceae Viscaceae Gymnosperms S Seed Conifers mostly herbaceous; without mycorrhiza Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae ANA grade G often unilocular with free-central placentation Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae pollen colpate, surface spiny Chenopodiaceae Droseraceae Nyctaginaceae Simmondsiaceae Magnoliids betalains or anthocyanins (latter, e.g., in Caryophyllaceae) aryophyllales S 37/749/11620 C Basellaceae Drosophyllaceae Phytolaccaceae Talinaceae

Angiosperms Monocots u mostly woody; lvs mostly undivided, hydathode teeth Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae p flw often 4-merous, K much smaller than C, persisting

Fabids e intrastaminal disk, G inferior; fr drupaceous Cornaceae Grubbiaceae Loasaceae diverse iridoids 6/51/590 Cornales alvids Curtisiaceae Hydrangeaceae Hydrostachyaceae Nyssaceae M r seed exo- a Actinidiaceae Ericaceae Pentaphylacaceae Sarraceniaceae Lamiids testal lvs teeth often theoid; nodes unilacunar s embryo flw 5-merous, pentacyclic Balsaminaceae Fouquieriaceae Polemoniaceae Styracaceae ampanulids C t long nonhydrolyzable tannins, ellagic acid, triterpenoids, hydroquinones Clethraceae Lecythidaceae Primulaceae Symplocaceae 22/346/11545 Ericales Diapensiaceae Marcgraviaceae Roridulaceae Theaceae e tropical trees; lvs exstipulate/entire; nodes often unilacunar Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae r pedicels articulated; style short; embryo long i endosperm copious indole and quinoline alkaloids (camptothecin) Oncothecaceae Icacinaceae d 2/24/202 Icacinales s tropical trees; few common features; nodes usu. trilacunar A Metteniusa: NE Andes, cloud montane ; lvs alternate; flw bisexual, fragrant; anthers sagittate Theodor C. H. Cole, Dipl. Biol. late pollen sacs moniliform, dehisce longitudinally; infl axillary cyme;G (5) unilocular Metteniusaceae s 1/11/55 Metteniusales Prof. Dr. sympetaly Hartmut H. Hilger, t woody; dioecious Dahlem Centre of Plant Sciences (DCPS) e flw small, C valvate, G unilocular; indehiscent iridoids (aucubin), gutta arryales Eucommiaceae Garryaceae (incl. Aucubaceae) Institute of Biology – Botany r L 2/3/18 G Freie Universität Berlin i a lvs spiral, simple; nodes unilacunar Altensteinstr. 6, D-14195 Berlin, Germany m flw petals plicate; K persisting Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae) d diverse alkaloids, no iridoids s ii 5/165/4125 Hydroleaceae Montiniaceae Sphenocleaceae d herbaceous; lvs opposite, glandular-pubescent mostly sympetalous s flw < 5 mm across, K valvate, G(3) inf, nectary C enclosing fr septicidal capsule, K/C/A persistent; seeds minute Vahliaceae Peter F. Stevens, Ph.D. A and G in bud Africa to India; iridoids 1/1/8 Vahliales Missouri Botanical Garden (MoBot) G(2) lvs opposite; interpetiolar stipules or nodal line/ridge style single, long C colleters Apocynaceae (incl. Asclepiadaceae) Loganiaceae St. Louis, MO 63166-0299, USA nectary gynoecial forming flw corolla convolute in bud A=C distinct entianales ovules unitegmic indole alkaloids, iridoids 5/1121/19915 Gentianaceae Gelsemiaceae Rubiaceae and polyandry tube G endosperm cellular rare lvs opposite; nodes 1:1; flw mostly monosymmetric University of Missouri – St. Louis iridoids + A A often 2(+2); gland-headed hairs with radially arranged cells Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae infl epipetalous Department of Biology cymose 6-oxygenated flavones, Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae St. Louis, MO 63121 - 4499, USA fr a drupe oligosaccharides cornoside, verbascoside (acetoside) amiales 24/1059/23810 L seed single Byblidaceae Martyniaceae Pedaliaceae Stilbaceae ellagic acid lvs roughly hairy; nodes unilacunar Calceolariaceae Gesneriaceae Oleaceae Phrymaceae Verbenaceae lacking infl scorpioid; mostly 4 ovules isokestose, higher inulins, Boraginaceae Codonaceae Coldeniaceae Cordiaceae Ehretiaceae (+ Lennoaceae) pyrrolizidine alkaloids 6-11/150/3095 Boraginales COLE TCH, HILGER HH, STEVENS PF (2019) Angiosperm Phylogeny Poster – Systematics Heliotropiaceae Hydrophyllaceae Namaceae Wellstediaceae woody; lvs serrate • hypothetical based on molecular phylogenetic data (Dec. 2018) flw 1-2 ovules/carpel, C± free, K slightly connate • phylogeny, classification, and features chiefly followAPweb and APG fr usually drupe with broad stigma quifoliales Aquifoliaceae Cardiopteridaceae Stemonuraceae • 64 orders and around 420 families currently recognized by APG IV (some minor families excluded here) early 5/21/536 A • contrary to APG, but in compliance with APweb and other seminal sources, we here recognize several families within Boraginales sympetaly herbs or ; flw often monosymmetric Asteraceae Goodeniaceae Pentaphragmataceae • branch lengths deliberate, not expressing actual time scale flw small plunger sec. pollination device (long style) • the characters listed do not necessarily apply to all members of a clade embryo nodes trilacunar Calyceraceae Menyanthaceae Rousseaceae short • position of various characters on the tree uncertain inulin, sesquiterpenes, secoiridoids Asterales 11/1743/26870 Campanulaceae (incl. Lobeliaceae) Stylidiaceae • for characteristics see: FGVP, Kubitzki K, ed. (1990 ff) • orders (and higher ranks) are linked to the according pages on APweb mostly woody; infl racemose, C free • numbers set in gray next to orders refer to families, genera, and species C anthers basifixed, nectary disc scalloniales Escalloniaceae 1/9/130 E * Ceratophyllum: the alleged tepals could be bracts (issue being addressed in ongoing research and debate); ** Rosids: phylogeny within the group controversial a This poster is now available in 24 languages (please refer to the authors’ pages on ResearchGate) woody; evergreen; nodes 1:1 m flw polysymmetric, anthers basifixed Bruniaceae Columelliaceae (incl. Desfontainia)

2/14/79 runiales References: Stevens PF (2019) APweb – www.mobot.org/MOBOT/research/APweb p B

) APG IV (2016); Judd W et al. (2016); Simpson M (2010); Soltis DE et al. (2005/2011/2014); Watson/Dallwitz (2018) woody (except Apiaceae) a lvs often divided; nodes usu. multilacunar Acknowledgements to our contributors, translators, and consultants: Rubina Abid, Julien B. Bachelier, Przemysław Baranow, Zoltán Barina, Fernanda Antunes Carvalho, Christoph Dobeš, Apiaceae Griseliniaceae Pennantiaceae n G inferior infl mostly umbel; drupe or schizocarp

CC-BY Ray F. Evert, Mohamed Fennane, Marc Gottschling, Zigmantas Gudžinskas, Akitoshi Iwamoto, Chen-Kun Jiang, Anna Kagiampaki, Kent Kainulainen, Elizabeth A. Kellogg, Sangtae Kim, ( (Apiaceae: mericarp/carpophore) Apiales Araliaceae Myodocarpaceae Pittosporaceae Aslı Doğru-Koca, Nikos Krigas, Sanjay Kumar, Diego Medan, Sergei L. Mosyakin, Sofi Mursidawati, Dashzeveg Nyambayar, Anastasiya V. Odintsova, Richard G. Olmstead, Batlai Oyuntsetseg, u fr usually 7/494/5489 Peter H. Raven, Yasaman Salmaki, Federico Selvi, Paramjit Singh, Douglas E. Soltis, Pramote Triboun, Zoya M. Tsymbalyuk, Magsar Urgamal, Maximilian Weigend, Michael Wink, Shahin Zarre 1–few-seeded l woody; infl racemose, flw 4-merous filaments stout; capsule septicidal Paracryphiaceae i 1/3/36 Paracryphiales Angiosperm Tracheophyte Bryophyte d lvs opposite, often basally connate Phylogeny Phylogeny Phylogeny s with scales flw often monosymmetric Adoxaceae Poster Poster Poster K persistent in fruit; secoiridoids ( ) 2/46/1090 ipsacales incl. Dipsacaceae Diervillaceae Linnaeaceae Morinaceae Valerianaceae © The Authors, 2019/1 © The D Caprifoliaceae