Angiosperm Phylogeny Flowering Plant Systematics

woody; vessels lacking

dioecious; flw T5–8, A∞, G5–8, 1 ovule/carpel, embryo sac 9-nucleate
A mborellales Amborellaceae

1/1/1
1 species, New Caledonia
A g aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A1–∞, embryo sac 4-nucleate
r seeds operculate with perisperm but endosperm reduced or small
N ymphaeales Cabombaceae Hydatellaceae Nymphaeaceae

3/6/74
N
mucilage; alkaloids (no benzylisoquinolines)
a
woody, vessels solitary
d
Austrobaileyales
ethereal oils in spherical idioblasts

flw T>10, A∞, G ca.9, embryo sac 4-nucleate

Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae

3/5/100
A e tiglic acid, aromatic terpenoids
lvs opposite, interpetiolar stipules
a flw small T0–3, A1–5, G1, 1 apical ovule/carpel
Chloranthales Chloranthaceae

1/4/75
e n nodes swollen
woody; foliar sclereids
a g
Canellales
sesquiterpenes

2/10/125
K and C distinct
aromatic terpenoids Canellaceae Winteraceae
r i
tension wood +
(pellucid dots)

nodes trilacunar ± herbaceous; lvs two-ranked, leaf base sheathing

l o
Piperales

4/17/4170
single adaxial prophyll
nodes swollen Aristolochiaceae (incl. Hydnoraceae) Piperaceae Saururaceae
y s woody; lvs opposite
Calycanthaceae Hernandiaceae Monimiaceae
Laurales

7/91/2858
flw with hypanthium, staminodes frequent
p anthers often valvate; carpels with 1 ovule; embryo large Gomortegaceae Lauraceae Siparunaceae
e Magnoliids woody; pith septate; lvs two-ranked
benzylisoquinoline alkaloids

Magnoliales Annonaceae Eupomatiaceae Magnoliaceae

6/128/3140
ovules with obturator
r features as in endosperm ruminate
Degeneriaceae Himantandraceae Myristicaceae
embryo sac 8-nucleate

m “Early Angiosperms” infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules

Acorales
endosperm triploid

ovules atropous, seeds with epidermal perisperm and copious endosperm

s Acoraceae

1/1/2-4
ethereal oils
mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules
Alismataceae Butomaceae Posidoniaceae Scheuchzeriaceae

14/166/4660
Alismatales
woody infl ± scapose; flw G apocarpous; placentation often laminar
vessels absent endosperm helobial; embryo large/green Aponogetonaceae Hydrocharitaceae Potamogetonaceae Tofieldiaceae
eustele
sieve tube plastids twining vines (often); lvs often reticulate
Araceae Juncaginaceae Ruppiaceae Zosteraceae

Dioscoreales

5/21/1050
with starch grains ovary often inferior, style short, branched
lvs simple, persistent, entire
flw strobilar, perfect, parts free Monocots Ca oxalate
steroidal sapogenins/alkaloids Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae Thismiaceae

Pandanales

5/36/1345
P parts varying, often in threes, raphides some woody (with terminally tufted lvs)
scattered bundles in stem
weakly differentiated
no secondary thickening endosperm infl sometimes with spathe + spadix Cyclanthaceae Pandanaceae Triuridaceae Velloziaceae
anthers tetrasporangiate nuclear
mostly herbaceous often geophytes (bulbs, tubers, rhizomes); leaf bases often not sheathing
stamen with broad filaments helobial
pollen monosulcate
Liliales
flw T sometimes spotted, nectaries on T, anthers extrorse
Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae

10/67/1558
pollen monosulcate
sieve tube plastids with seeds many
G apocarpous (style short in most)
compitum (if present) extragynoecial
protein crystals
sympodial branching
phytomelan lacking; fructans in stems, chelidonic acid Colchicaceae Liliaceae Petermanniaceae Smilacaceae
nectaries absent stem with ring of bundles

Petrosaviales
lvs parallel-veined, entire
extrafloral nectaries

siphonogamy fr follicle
Petrosaviaceae

1/2/3
no glandular teeth East Asia
double fertilization > endosperm
flw pentacyclic
embryo very small

14/1122/36205
P 3-merous, A opp. P often geophytes Amaryllidaceae (incl. Agapanthaceae, Alliaceae) Hypoxidaceae Iridaceae
filaments narrow
anthers broadly attached
septal nectary
fr capsule or berry
seed coat obliterated or with phytomelan Asparagales Asparagaceae (incl. Agavaceae, Hyacinthaceae, Ruscaceae) Lanariaceae Orchidaceae
Tecophilaeaceae Asphodelaceae (incl. Xanthorrhoeaceae, Hemerocallidaceae)
woody, often monopodial
single cotyledon

Arecales

1/188/2585
lvs often palmately or pinnately pseudocompound, reduplicate-plicate
radicle not persistent
stem-borne roots numerous
intense primary growth, large apical meristem, infl often with spathe
alkaloids
Arecaceae Dasypogonaceae
Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae

15/997/18875
mostly herbaceous; lvs grassy; epidermis siliceous
mycorrhiza – (in most)
flw often anemophilous, minute, chaffy, without nectaries Poales Cyperaceae Juncaceae Rapateaceae Typhaceae (incl. Sparganiaceae)

Commelinids
flw monosymmetric or not, few fertile stamens
Commelinaceae Haemodoraceae Hanguanaceae
infl thyrsus of scorpioid cymes
Commelinales

5/68/812
phenylphenalenones
Philydraceae Pontederiaceae
UV-fluorescing cell walls rhizomatous, large-leafed herbs; pseudostem common
(ferulic/coumaric acids) flw irregular/monosymmetric, septal nectaries
Cannaceae Heliconiaceae Marantaceae Strelitziaceae
Zingiberales

8/92/~2500
silicic acid in leaves A often strongly modified/reduced, G inferior; seeds often arillate
cuticular waxes often in rodlets
aggregated into scallops
silicic acid Costaceae Lowiaceae Musaceae Zingiberaceae
aquatic; herbaceous; monoecious

Ceratophyllales
lvs whorled, no pellucid dots; vessels lacking
flw T0 or 9–10*, A1, G1, 1 apical ovule/carpel
Ceratophyllaceae

1/1/6
pollen inaperturate, pollen tube branched, hydrophilous
lvs often divided
Berberidaceae Eupteleaceae Menispermaceae
Ranunculales

7/199/4510
flw parts whorled, P single or multiple whorls
G apocarpous/paracarpous, superior Ranunculaceae
ethereal berberines Circaeasteraceae Lardizabalaceae Papaveraceae
oils

Proteales
mostly woody

4/85/1750
not in
idioblasts
flw T often 4-merous
A epitepalous, connectives sometimes with apical appendage
Nelumbonaceae Platanaceae Proteaceae Sabiaceae

Trochodendrales
woody; vessels lacking
flw T missing, A∞, G>5 laterally connate with abaxial nectaries Trochodendraceae

1/2/2
fr aggregate of follicles

E
recepta-
cular

Buxales
nectary mostly woody; lvs evergreen, stomata cyclocytic
Buxaceae (incl. Haptanthaceae)

1/7/120
u common mostly monoecious, flw unisexual, T ± uniform or missing
pregnane pseudoalkaloids
d
Gunnerales
lvs toothed, sec. veins palmate
i Gunneraceae Myrothamnaceae

2/2/50
dioecious, flw unisexual, T small to lacking
c
absent

ellagic acid

o
Dilleniales
mostly woody; lvs if veins strong, proceed to apex of teeth
t flw mostly K5, persisting, mostly A∞, G mostly slightly connate Dilleniaceae

1/10/300
benzylisoquinolines

seeds often with aril; fr usually follicles

s Altingiaceae Cynomoriaceae Haloragaceae Peridiscaceae
lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent
pollen tricolpate
protandry common
fr mostly dry, dehiscent
myricetin, flavonols
15/112/2500
Saxifragales Cercidiphyllaceae Daphniphyllaceae Hamamelidaceae Paeoniaceae
flw K/C/P opp A
often tendrillar vines; lvs often divided and with glandular teeth
Crassulaceae Grossulariaceae Iteaceae Saxifragaceae
filaments rather narrow
A epipetalous, 2 ovules per carpel; raphides, pearl glands
Vitales
1/14/850

nodes 3:3
stomata anomocytic berries Vitaceae
microsporogenesis
simultaneous cork origin deep-seated
endosperm lacking
Zygophyllales
2/24/345

resinous, lignans/neolignans, harman alkaloids Krameriaceae Zygophyllaceae

ellagic and gallic acids

lvs often compound, pulvini (sleep movement)

stipules
Brunelliaceae Connaraceae Elaeocarpaceae
Oxalidales
flw A5 or multiple, branched style common
7/60/1845

mucilage cells Oxalidaceae

Cephalotaceae Cunoniaceae Huaceae
S
oxalates

infl cymose, flw small

u
Celastrales
2/94/1355

G often 3-merous, nectary often intrastaminal disk

COM clade Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae) Lepidobotryaceae
p seeds often arillate (red-orange) or winged

e Achariaceae Euphorbiaceae Rafflesiaceae Ochnaceae Podostemaceae

r F habits and habitats extremely diverse
36/716/16065

c E r a
lvs margins toothed
flw G often tricarpellate Malpighiales Chrysobalanaceae
Clusiaceae
Hypericaceae
Linaceae
Passifloraceae
Phyllanthaceae
Rhizophoraceae
Salicaceae
b
o u o Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae
R
flw often “papilionaceous”: wing, standard, keel, C clawed, G1 (most), A10 (most)
4/754/20140

d s
r
e
i i
c d
i fr pod
o d
symbiosis with root nodule bacteria
diverse alkaloids, NP amino acids, lectins (in Fabaceae) Fabales Fabaceae Polygalaceae Quillajaceae Surianaceae
s s lvs mostly simple with stipules
Barbeyaceae Elaeagnaceae Rosaceae
i flw K valvate (and hypanthium) persisting
9/261/7725

o
t
s
d N fix
carpels with 1 ovule, stigma dry
dihydroflavonols Rosales Cannabaceae
Dirachmaceae
Moraceae
Rhamnaceae
Ulmaceae
Urticaceae (incl. Cecropiaceae)
s s embryo large
endosperm scanty
lvs mostly alternate
flw often unisexual, G mostly inferior

Cucurbitales
7/109/2935

parietal placentation Apodanthaceae Begoniaceae Corynocarpaceae Datiscaceae

cucurbitacins
Anisophyllaceae Coriariaceae Cucurbitaceae Tetramelaceae
mostly trees; lvs mostly undivided; flw small, unisexual
anemophilous, thus T reduced or lacking, G mostly inferior
Betulaceae Fagaceae Myricaceae
Fagales
7/33/1005

infl spikes or catkins; fr 1-seeded, mostly nuts

Ticodendraceae
ectomycorrhiza; tannins, dihydroflavonols Casuarinaceae Juglandaceae Nothofagaceae
stems sometimes jointed at nodes; lvs with glandular teeth
2(5)/17/897

flw A obdiplostemonous, nectary outside A; fr capsule

ethereal oils, ellagic acid Geraniales Geraniaceae Francoaceae (incl. Ledocarpaceae, Melianthaceae, Vivianiaceae)
lvs opposite, colleters (glandular hair on adaxial surface of petiole base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae)
stipules small (if any), cork deep seated
9/380/13005

Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae)

Myrtales
flw 5-merous flw K valvate, persisting, A incurved in bud, ovary inferior, ovules many
parts whorled
K + C free P endosperm scanty, scaly bark, flavonols, myricetin
Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae
diplostemonous e woody
Crossosomataceae Stachyuraceae
n Crossosomatales
pollen tricolporate stylodia free
7/12/66

G connate hypanthium with nectary Strasburgeriaceae

style + t Geissolomataceae Staphyleaceae
endosperm nuclear
a M trees, dioecious, lvs compound

Picramniales
extrafloral nectaries, staminate flw: A = and opposite C
1/3/49

p a bark bitter, anthraquinones Picramniaceae

e l mostly woody (silica/silicified)
t v Anacardiaceae Burseraceae Kirkiaceae Nitrariaceae Sapindaceae
9/479/6550

lvs often alternate, odd-pinnately compound

a i
flw often imperfect, intrastaminal disk
ethereal oils Sapindales Biebersteiniaceae Meliaceae Rutaceae Simaroubaceae
l d
a s
vessel elements with scalariform perforations; mucilage cells
embryo lvs margins toothed, stipules cauline
Huerteales Dipentodontaceae Gerrardinaceae Petenaeaceae Tapisciaceae
4/6/24

e large flw small, A = and opposite K, ovules 1-2/carpel

endo-
sperm bark fibrous; hairs often stellate Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
10/338/6005

scanty
Liverworts

Mosses
flw K often valvate, petals contorted, A often ∞; mucilage
cyclopropenoid fatty acids, flavones Malvales Cistaceae
Dipterocarpaceae
Cytinaceae
Neuradaceae
Muntingiaceae
Sphaerosepalaceae
Sarcolaenaceae
Thymelaeaceae
woody or herbs
Bataceae Caricaceae Limnanthaceae Salvadoraceae
18/405/5035

Brassicales
Hornworts flw often 4-merous, often clawed petals, infl racemose

Lycophytes
myrosin cells, glucosinolates
Brassicaceae Cleomaceae Moringaceae Tovariaceae
woody; (semi-)parasites; mycorrhiza absent; lvs margins entire Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae
Ferns
13/151/1992

flw A epipetalous; perianth often simple, valvate, persisting

Tracheophytes
Santalales Balanophoraceae Misodendraceae Opiliaceae Schoepfiaceae
(incl. horsetails)
carpels/ovaries/ovules often reduced, placentation free-central; fr drupe, one-seeded
Cycads polyacetylenes, triterpene sapogenins, silicic acid
Ginkgo Loranthaceae Olacaceae Santalaceae Viscaceae
Ephedra woody
lvs stomata cyclocytic, petiole bundles annular

Berberidopsidales
Welwitschia
2/3/4

fr indehiscent (more or less fleshy)

Seed Gymnosperms Gnetum
Conifers calcium oxalate as crystals Aextoxicaceae Berberidopsidaceae
Plants
ANA grade Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae
mostly herbaceous; without mycorrhiza
37/749/11620

Angiosperms Magnoliids G often unilocular with free-central placentation Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae

Monocots
S pollen colpate, surface spiny
betalains or anthocyanins (latter, e.g., in Caryophyllaceae) Caryophyllales Chenopodiaceae
Basellaceae
Droseraceae
Drosophyllaceae
Nyctaginaceae
Phytolaccaceae


Simmondsiaceae
Talinaceae
u Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae
Fabids p mostly woody; lvs mostly undivided, hydathode teeth
flw often 4-merous, K much smaller than C, persisting Cornaceae Grubbiaceae Loasaceae
Cornales
6/51/590

Malvids e intrastaminal disk, G inferior; fr drupaceous

Rosids
r
diverse iridoids Curtisiaceae Hydrangeaceae Hydrostachyaceae Nyssaceae
Lamiids seed
Asterids a exo- lvs teeth often theoid; nodes unilacunar Actinidiaceae Ericaceae Pentaphylacaceae Sarraceniaceae
22/346/11545

testal flw 5-merous, pentacyclic

s Balsaminaceae Fouquieriaceae Polemoniaceae Styracaceae
Ericales
Campanulids nonhydrolyzable tannins, ellagic acid
embryo Clethraceae Lecythidaceae Primulaceae Symplocaceae
t long
triterpenoids, hydroquinones
Diapensiaceae Marcgraviaceae Roridulaceae Theaceae
e trop. trees; lvs exstipulate/entire; nodes often unilacunar Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae
r pedicels articulated; style short; embryo long

Icacinales
2/24/202

endosperm copious
i indole and quinoline alkaloids (camptothecin) Oncothecaceae Icacinaceae
d trop. trees; few shared features; nodes usu. trilacunar
alkaloids, iridoids
Theodor C. H. Cole, Dipl. Biol. s
A Metteniusales Metteniusaceae
1/11/55

Metteniusa: lvs alternate; flw bisexual, fragrant; anthers sagittate; pollen sacs moniliform, dehisce longitudinally
late infl axillary cyme; G(5) unilocular; NE Andes, cloud montane forest
Hartmut H. Hilger, Prof. Dr. s sympetaly
woody; dioecious
Dahlem Centre of Plant Sciences (DCPS) t flw small, C valvate, G unilocular
Institute of Biology – Botany e L
fr indehiscent
Garryales Eucommiaceae Garryaceae (incl. Aucubaceae)
2/3/18

iridoids (aucubin), gutta

Freie Universität Berlin r a lvs opposite; nodes 1:1 Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae
24/1059/23810

Altensteinstr. 6, D-14195 Berlin, Germany i m flw mostly monosymmetric

d ii
A often 2(+2); gland-headed hairs with radially arranged cells
6-oxygenated flavones, oligosaccharides cornoside, verbascoside (acetoside) Lamiales Bignoniaceae
Byblidaceae
Lentibulariaceae
Martyniaceae
Paulowniaceae
Pedaliaceae
Scrophulariaceae
Stilbaceae
s d herbaceous; lvs opposite, glandular-pubescent Calceolariaceae Gesneriaceae Oleaceae Phrymaceae Verbenaceae
s flw < 5 mm across, K valvate, G(3) inf, nectary
Peter F. Stevens, Ph.D. mostly sympetalous
fr septicidal capsule, K/C/A persistent; seeds minute
Vahliales Vahliaceae
1/1/8

Africa to India; iridoids

C enclosing
Missouri Botanical Garden (MoBot) A and G in bud lvs spiral, simple; nodes unilacunar
St. Louis, MO 63166-0299, USA G(2) flw petals plicate
5/165/4125

Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae)

Solanales
style single, long C K persistent in fruit
and nectary gynoecial A=C forming
alkaloids diverse, no iridoids
University of Missouri – St. Louis ovules unitegmic polyandry distinct
tube
Hydroleaceae Montiniaceae Sphenocleaceae
endosperm cellular rare lvs opposite
Department of Biology
5/1121/19915

iridoids + infl A interpetiolar stipules or nodal line/ridge, colleters

Gentianales Apocynaceae (incl. Asclepiadaceae) Loganiaceae

epipetalous
St. Louis, MO 63121 - 4499, USA cymose flw corolla convolute in bud
fr drupe indole alkaloids, iridoids
seed single Gentianaceae Gelsemiaceae Rubiaceae
ellagic acid lvs roughly hairy; nodes unilacunar
6-11/150/3095

infl scorpioid
lacking
Boraginaceae Codonaceae Coldeniaceae Cordiaceae Ehretiaceae (+ Lennoaceae)
COLE TCH, HILGER HH, STEVENS PF (2019) Angiosperm Phylogeny Poster – Flowering Plant Systematics
mostly 4 ovules
isokestose, higher inulins, pyrrolizidine alkaloids Boraginales Heliotropiaceae Hydrophyllaceae Namaceae Wellstediaceae
• hypothetical tree based on molecular phylogenetic (chiefly plastid) data (May 2019) woody; lvs serrate
• phylogeny, classification, and features chiefly follow APweb, APG, and PPA (Plastid Phylogenomic Angiosperm see Li HT et al. 2019) flw 1-2 ovules/carpel, C ± free, K slightly connate
Aquifoliales Aquifoliaceae Cardiopteridaceae Stemonuraceae
5/21/536

fr usu. drupe with broad stigma

• 64 orders and around 420 families currently recognized by APG IV (some minor families omitted here)
• contrary to APG, but in compliance with APweb and other seminal sources, we here recognize several families within Boraginales herbs or shrubs; nodes trilacunar
Asteraceae Goodeniaceae Pentaphragmataceae
11/1743/26870

• branch lengths deliberate, not expressing actual time scale early flw often monosymmetric
• the characters listed do not necessarily apply to all members of a clade
• position of various characters on the tree uncertain
sympetaly
flw small
embryo
plunger sec. pollination device (long style)
inulin, sesquiterpenes, secoiridoids Asterales Calyceraceae Menyanthaceae Rousseaceae
• for family characteristics see: FGVP, Kubitzki K, ed. (1990 ff) short
mostly woody
Campanulaceae (incl. Lobeliaceae) Stylidiaceae
• orders (and higher ranks) are linked to the according pages on APweb
Escalloniales
infl racemose, C free
1/9/130

• numbers set in gray next to orders refer to families, genera, and species Escalloniaceae
C
anthers basifixed, nectary disk
* Ceratophyllum: the alleged tepals could be bracts (issue being addressed in ongoing research and debate)
This poster is now available in 25 languages (please refer to the authors’ pages on ResearchGate) a woody; evergreen

Bruniales
2/14/79

nodes 1:1
References: Stevens PF (2019) APweb – www.mobot.org/MOBOT/research/APweb
m flw polysymmetric, anthers basifixed 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)
p woody (except Apiaceae)
© The Authors, 2019/3 (CC-BY)

Acknowledgements to our contributors, translators, and consultants: Rubina Abid, Julien B. Bachelier, Przemysław Baranow, Zoltán Barina, Fernanda Antunes Carvalho, Christoph Dobeš,
a lvs often divided; nodes usu. multilacunar
Apiaceae Griseliniaceae Pennantiaceae
7/494/5489

A piales
Ray F. Evert, Mohamed Fennane, Marc Gottschling, Zigmantas Gudžinskas, Akitoshi Iwamoto, Chen-Kun Jiang, Anna Kagiampaki, Kent Kainulainen, Elizabeth A. Kellogg, Sangtae Kim, G
infl mostly umbel; fr drupe or schizocarp
Aslı Doğru-Koca, Nikos Krigas, Sanjay Kumar, Diego Medan, Sergei L. Mosyakin, Sofi Mursidawati, Dashzeveg Nyambayar, Anastasiya V. Odintsova, Richard G. Olmstead, Batlai Oyuntsetseg,
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 inferior
fr
(Apiaceae: mericarp/carpophore) Araliaceae Myodocarpaceae Pittosporaceae
u usually woody
l
Paracryphiales
1–few-seeded
infl racemose, flw 4-merous, filaments stout
1/3/36

capsule septicidal Paracryphiaceae

Angiosperm Tracheophyte Bryophyte i lvs opposite, often basally connate
Phylogeny Phylogeny Phylogeny d buds with scales; flw often monosymmetric Adoxaceae
Dipsacales
2/46/1090

Poster Poster Poster s K persistent in fruit

secoiridoids Caprifoliaceae (incl. Dipsacaceae Diervillaceae Linnaeaceae Morinaceae Valerianaceae)

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