Angiosperm Phylogeny

Flowering Plant Systematics

A
woody, vessels lacking; dioecious; flw T5–8, A∞, G5–8, 1 ovule/carpel, embryo sac 9-nucleate; 1 species, New Caledonia A mborellales Amborellaceae

1/1/1
g

N ymphaeales
aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A1–∞, embryo sac 4-nucleate
r
Cabombaceae Hydatellaceae Nymphaeaceae
N
seeds operculate with perisperm but endosperm reduced or small; mucilage; alkaloids (no benzylisoquinolines)

3/6/74
a
woody, vessels solitary; flw T>10, A∞, G ca.9, embryo sac 4-nucleate
Austrobaileyales
ethereal oils in spherical idioblasts

d
Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae

3/5/100
A
tiglic acid, aromatic terpenoids
e

a Chloranthales
lvs opposite, interpetiolar stipules; nodes swollen
flw small T0–3, A1–5, G1, 1 apical ovule/carpel Chloranthaceae
e n

1/4/75
a g
Canellales
woody; foliar sclereids, K and C distinct
sesquiterpenes

Canellaceae Winteraceae

2/10/125
aromatic terpenoids
r i
tension wood +
(pellucid dots)

nodes trilacunar
l o
Piperales
± herbaceous; lvs two-ranked, leaf base sheathing
Aristolochiaceae (incl. Hydnoraceae) Piperaceae Saururaceae

4/17/4170
single adaxial prophyll; swollen nodes
y s
Calycanthaceae Hernandiaceae Monimiaceae
Laurales
woody; lvs opposite; flw with hypanthium, staminodes frequent

7/91/2858
p often valvate anthers; carpels with 1 ovule; embryo large
Gomortegaceae Lauraceae Siparunaceae
e Magnoliids
benzylisoquinoline alkaloids

Magnoliales Annonaceae Eupomatiaceae Magnoliaceae

woody; pith septate; lvs two-ranked; ovules with obturator

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

m “Early Angiosperms”

Acorales
infl spadix with spathe; lvs axils with mucilaginous intravaginal squamules
endosperm triploid

s ovules atropous, seeds with epidermal perisperm and copious endosperm; ethereal oils Acoraceae

1/1/2-4
Alismataceae Butomaceae Posidoniaceae Scheuchzeriaceae
Alismatales

14/166/4660
woody; vessels absent mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules
eustele infl ± scapose; flw G apocarpous; placentation often laminar; endosperm helobial; embryo large/green Aponogetonaceae Hydrocharitaceae Potamogetonaceae Tofieldiaceae
sieve tube plastids Araceae Juncaginaceae Ruppiaceae Zosteraceae
with starch grains stem with ring of bundles
lvs simple, persistent, entire
Monocots Ca oxalate
fr a follicle
East Asia Petrosaviales Petrosaviaceae

1/2/3
flw strobilar, perfect, parts free raphides
P parts varying, often in threes,
weakly differentiated scattered bundles in stem endosperm often twining vines; lvs often reticulate
Dioscoreales Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae Thismiaceae

5/21/1050
anthers tetrasporangiate nuclear ovary often inferior, style short, branched; steroidal sapogenins/alkaloids
no secondary thickening helobial
stamen with broad filaments mostly herbaceous
pollen monosulcate pollen monosulcate some woody (with terminally tufted leaves)
Pandanales Cyclanthaceae Pandanaceae Triuridaceae Velloziaceae

5/36/1345
G apocarpous (style short in most) sieve tube plastids with infl sometimes spathe + spadix
compitum (if present) extragynoecial protein crystals
often geophytes (bulbs, tubers, rhizomes); leaf bases often not sheathing
nectaries absent sympodial branching Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae
Liliales

10/67/1558
siphonogamy lvs parallel-veined, entire flw tepals sometimes spotted, nectaries on tepals, anthers extrorse
extrafloral nectaries

double fertilization > endosperm no glandular teeth many seeds; phytomelan lacking; fructans in stems, chelidonic acid Colchicaceae Liliaceae Petermanniaceae Smilacaceae
embryo very small flw pentacyclic
P 3-merous, A opp. P often geophytes
Amaryllidaceae ( incl. Agapanthaceae, Alliaceae) Hypoxidaceae Iridaceae

14/1122/36205
Asparagales
filaments narrow capsule or berry
anthers broadly attached seed coat obliterated or with phytomelan Asparagaceae ( incl. Agavaceae, Hyacinthaceae, Ruscaceae) Lanariaceae Orchidaceae
septal nectary
single cotyledon Tecophilaeaceae Asphodelaceae ( incl. Xanthorrhoeaceae, Hemerocallidaceae)
radicle not persistent
woody, often monopodial

1/188/2585
Arecales
stem-borne roots numerous lvs often palmately or pinnately pseudocompound, reduplicate-plicate
intense primary growth, large apical meristem, infl often with spathe; alkaloids Arecaceae Dasypogonaceae

15/997/18875
Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae
Poales
mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent
lvs grassy; flw often anemophilous, minute, chaffy, without nectaries
Cyperaceae Juncaceae Rapateaceae Typhaceae (incl. Sparganiaceae)
flw monosymmetric or not, few fertile stamens
Commelinids infl thyrsus of scorpioid cymes
Commelinales Commelinaceae Haemodoraceae Hanguanaceae

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
silicic acid in leaves

8/92/~2500
A often strongly modified/reduced, G inferior
cuticular waxes often in rodlets
aggregated into scallops
seeds often arillate, silicic acid 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
pollen inaperturate, pollen tube branched, hydrophilous Ceratophyllales Ceratophyllaceae

1/1/6
Berberidaceae Eupteleaceae Menispermaceae
Ranunculales
lvs often divided; flw parts whorled, P single or multiple whorls

7/199/4510
G apocarpous/paracarpous, superior; berberines Ranunculaceae
ethereal
oils
Circaeasteraceae Lardizabalaceae Papaveraceae

Proteales
mostly woody; flw tepals often 4-merous

4/85/1750
not in
idioblasts A epitepalous, connectives sometimes with apical appendage Nelumbonaceae Platanaceae Proteaceae Sabiaceae
woody; vessels lacking; flw tepals missing, A∞
G>5 laterally connate with abaxial nectaries; fr aggregate of follicles Trochodendrales Trochodendraceae

1/2/2
E
recepta-
cular

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

1/7/120
common flw tepals ± uniform or missing; pregnane pseudoalkaloids

d dioecious, flw unisexual; lvs toothed, sec. veins palmate

i flw tepals small to lacking
ellagic acid Gunnerales Gunneraceae Myrothamnaceae

2/2/50
c
absent

o mostly woody; lvs if veins strong, proceed to apex of teeth

t
flw mostly K5, persisting, mostly A∞, G mostly slightly connate
Dilleniales Dilleniaceae

1/10/300
seeds often with aril; fr usually follicles
benzylisoquinolines

s lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent Altingiaceae Cynomoriaceae Haloragaceae Peridiscaceae
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

Vitales
nodes 3:3 A epipetalous, 2 ovules per carpel; raphides, pearl glands
1/14/850

stomata anomocytic berries Vitaceae

microsporogenesis
simultaneous cork origin deep-seated
endosperm lacking
Zygophyllales Krameriaceae Zygophyllaceae
2/24/345

resinous, lignans/neolignans, harman alkaloids

ellagic and gallic acids

stipules infl cymose, flw small

G often 3-merous, nectary often intrastaminal disk
Celastrales Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae) Lepidobotryaceae
2/94/1355

seeds often arillate (red-orange) or winged

S
u lvs often compound, pulvini (sleep movement)
Brunelliaceae Connaraceae Elaeocarpaceae
p
COM clade flw A5 or multiple, branched style common
Oxalidales Oxalidaceae
7/60/1845

mucilage cells; oxalates Cephalotaceae Cunoniaceae Huaceae

e
r F habits and habitats extremely diverse Achariaceae Euphorbiaceae Rafflesiaceae Ochnaceae Podostemaceae
E Malpighiales
36/716/16065

r a
lvs margins toothed Chrysobalanaceae Hypericaceae Passifloraceae Rhizophoraceae
c flw G often tricarpellate
Clusiaceae Linaceae Phyllanthaceae Salicaceae
o u o
R b
Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae
r d s i flw often “papilionaceous”: wing, standard, keel, C clawed, mostly G1
o
Fabales
4/754/20140

mostly A10; fr a pod; symbiosis with root nodule bacteria

i i d Fabaceae Polygalaceae Quillajaceae Surianaceae
e c d s s
diverse alkaloids, NP amino acids, lectins (in Fabaceae)

o s i lvs mostly simple with stipules Barbeyaceae Elaeagnaceae Rosaceae

t d flw K valvate (and hypanthium) persisting
Rosales
9/261/7725

N fix Cannabaceae Moraceae Ulmaceae

s s carpels with 1 ovule, stigma dry; dihydroflavonols
Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae)
embryo large
endosperm scanty lvs mostly alternate

Cucurbitales Apodanthaceae Begoniaceae Corynocarpaceae Datiscaceae

flw often unisexual, G mostly inferior
7/109/2935

parietal placentation; 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
infl spikes or catkins; fr 1-seeded, mostly nuts
Fagales
7/33/1005

Ticodendraceae
ectomycorrhiza; tannins, dihydroflavonols Casuarinaceae Juglandaceae Nothofagaceae
stems sometimes jointed at nodes; lvs with glandular teeth

Geraniales Geraniaceae Francoaceae (incl. Ledocarpaceae, Melianthaceae, Vivianiaceae)

2(5)/17/897

flw A obdiplostemonous, nectary outside A; fr capsule

ethereal oils, ellagic acid
flw 5-merous lvs opposite, colleters (glandular hair on adaxial surface of petiole base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae)
parts whorled
P stipules small (if any), cork deep seated

Myrtales
Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae)
9/380/13005

K + C free flw K valvate, persisting, A incurved in bud, ovary inferior, ovules many
diplostemonous e endosperm scanty, scaly bark, flavonols, myricetin Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae
pollen tricolporate
n woody

Crossosomatales
G connate
t stylodia free Crossosomataceae Stachyuraceae
style + Strasburgeriaceae
7/12/66

hypanthium with nectary

endosperm nuclear
a Geissolomataceae Staphyleaceae
M
trees, dioecious, lvs compound, extrafloral nectaries
p
e
staminate flw: A = and opposite C
Picramniales Picramniaceae
1/3/49

bark bitter, anthraquinones

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

lvs often alternate, odd-pinnately compound

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

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

s large
Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
endo- bark fibrous; hairs often stellate

Malvales
sperm
Cistaceae Cytinaceae Muntingiaceae Sarcolaenaceae
10/338/6005

scanty flw K often valvate, petals contorted, A often ∞; mucilage

Liverworts cyclopropenoid fatty acids, flavones
Dipterocarpaceae Neuradaceae Sphaerosepalaceae Thymelaeaceae
Mosses
woody or herbs; flw often 4-merous Bataceae Caricaceae Limnanthaceae Salvadoraceae
Brassicales
18/405/5035

Hornworts often clawed petals, infl racemose

myrosin cells, glucosinolates Brassicaceae Cleomaceae Moringaceae Tovariaceae
Lycophytes
woody; lvs stomata cyclocytic
Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae

Berberidopsidales
Ferns petiole bundles annular; fr indehiscent, more or less fleshy
Tracheophytes (incl. horsetails)
calcium oxalate as crystals Aextoxicaceae Berberidopsidaceae
2/3/4

Cycads
Ginkgo woody; (semi-)parasites; mycorrhiza absent; lvs margins entire
flw A epipetalous; perianth often simple, valvate, persisting Balanophoraceae Misodendraceae Opiliaceae Schoepfiaceae
Santalales
13/151/1992

Ephedra
carpels/ovaries/ovules often reduced, placentation free-central; fr drupe, one-seeded
Welwitschia
Gnetum polyacetylenes, triterpene sapogenins, silicic acid Loranthaceae Olacaceae Santalaceae Viscaceae
Seed Gymnosperms Conifers
Plants mostly herbaceous; without mycorrhiza Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae
ANA grade G often unilocular with free-central placentation Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae
Caryophyllales
37/749/11620

Magnoliids S pollen colpate, surface spiny

betalains or anthocyanins (latter, e.g., in Caryophyllaceae)
Chenopodiaceae
Basellaceae
Droseraceae
Drosophyllaceae
Nyctaginaceae
Phytolaccaceae


Simmondsiaceae
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
Cornales Cornaceae Grubbiaceae Loasaceae
6/51/590

Malvids r seed
diverse iridoids Curtisiaceae Hydrangeaceae Hydrostachyaceae Nyssaceae
Rosids

Lamiids
a exo-
testal Actinidiaceae Ericaceae Pentaphylacaceae Sarraceniaceae
s lvs teeth often theoid; nodes unilacunar
Balsaminaceae Fouquieriaceae Polemoniaceae Styracaceae
22/346/11545

Asterids Campanulids t
embryo
long
flw 5-merous, pentacyclic
nonhydrolyzable tannins, ellagic acid, triterpenoids, hydroquinones Ericales Clethraceae Lecythidaceae Primulaceae Symplocaceae
e tropical trees; lvs exstipulate/entire; nodes often unilacunar
Diapensiaceae Marcgraviaceae Roridulaceae Theaceae
r pedicels articulated; style short; embryo long
Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae
i endosperm copious
Icacinales
2/24/202

indole and quinoline alkaloids (camptothecin) Oncothecaceae Icacinaceae

d
s A tropical trees; few common features; nodes usu. trilacunar

Theodor C. H. Cole, Dipl. Biol. s

Metteniusa: NE Andes, cloud montane forest; lvs alternate; flw bisexual, fragrant; anthers sagittate
Metteniusales Metteniusaceae
1/11/55

late pollen sacs moniliform, dehisce longitudinally; infl axillary cyme; G(5) unilocular
sympetaly
Hartmut H. Hilger, Prof. Dr. t woody; dioecious
Dahlem Centre of Plant Sciences (DCPS)
Institute of Biology – Botany
e
r L
flw small, C valvate, G unilocular; fruit indehiscent
iridoids (aucubin), gutta Garryales Eucommiaceae Garryaceae (incl. Aucubaceae)
2/3/18

Freie Universität Berlin i a lvs spiral, simple; nodes unilacunar

m Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae)
Altensteinstr. 6, D-14195 Berlin, Germany d flw petals plicate; K persisting
Solanales
5/165/4125

s ii diverse alkaloids, no iridoids

Hydroleaceae Montiniaceae Sphenocleaceae
d herbaceous; lvs opposite, glandular-pubescent
flw < 5 mm across, K valvate, G(3) inf, nectary
s
Vahliales
mostly sympetalous

Peter F. Stevens, Ph.D.

C enclosing fr septicidal capsule, K/C/A persistent; seeds minute
Vahliaceae
1/1/8

A and G in bud Africa to India; iridoids

Missouri Botanical Garden (MoBot) G(2) lvs opposite; interpetiolar stipules or nodal line/ridge
C Apocynaceae (incl. Asclepiadaceae) Loganiaceae
Gentianales
style single, long
5/1121/19915

colleters
St. Louis, MO 63166-0299, USA nectary gynoecial forming flw corolla convolute in bud
and ovules unitegmic A=C distinct indole alkaloids, iridoids Gentianaceae Gelsemiaceae Rubiaceae
polyandry tube
endosperm cellular rare
University of Missouri – St. Louis lvs opposite; nodes 1:1; flw mostly monosymmetric
iridoids + A A often 2(+2); gland-headed hairs with radially arranged cells
Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae
24/1059/23810

Lamiales
infl
Department of Biology cymose epipetalous
6-oxygenated flavones, Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae
St. Louis, MO 63121 - 4499, USA fr a drupe oligosaccharides cornoside, verbascoside (acetoside) Byblidaceae Martyniaceae Pedaliaceae Stilbaceae
seed single
ellagic acid
lacking
lvs roughly hairy; nodes unilacunar Calceolariaceae Gesneriaceae Oleaceae Phrymaceae Verbenaceae
infl scorpioid; mostly 4 ovules
6-11/150/3095

isokestose, higher inulins,

pyrrolizidine alkaloids Boraginales Boraginaceae Codonaceae Coldeniaceae Cordiaceae Ehretiaceae (+ Lennoaceae)
Heliotropiaceae Hydrophyllaceae Namaceae Wellstediaceae
COLE TCH, HILGER HH, STEVENS PF (2019) Angiosperm Phylogeny Poster – Flowering Plant Systematics
woody; lvs serrate
• hypothetical tree based on molecular phylogenetic data (Dec. 2018)
• phylogeny, classification, and features chiefly follow APweb and APG
flw 1-2 ovules/carpel, C± free, K slightly connate
Aquifoliales Aquifoliaceae Cardiopteridaceae Stemonuraceae
5/21/536

fr usually drupe with broad stigma

• 64 orders and around 420 families currently recognized by APG IV (some minor families excluded here) early
• contrary to APG, but in compliance with APweb and other seminal sources, we here recognize several families within Boraginales sympetaly herbs or shrubs; flw often monosymmetric Asteraceae Goodeniaceae Pentaphragmataceae
• branch lengths deliberate, not expressing actual time scale flw small

Asterales
plunger sec. pollination device (long style)
11/1743/26870

• the characters listed do not necessarily apply to all members of a clade embryo nodes trilacunar Calyceraceae Menyanthaceae Rousseaceae
• position of various characters on the tree uncertain short
inulin, sesquiterpenes, secoiridoids Campanulaceae (incl. Lobeliaceae) Stylidiaceae
• for family characteristics see: FGVP, Kubitzki K, ed. (1990 ff)
• orders (and higher ranks) are linked to the according pages on APweb
C mostly woody; infl racemose, C free
Escalloniales Escalloniaceae
1/9/130

• numbers set in gray next to orders refer to families, genera, and species anthers basifixed, nectary disc
* 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) m woody; evergreen; nodes 1:1
Bruniales Bruniaceae Columelliaceae (incl. Desfontainia)
2/14/79

flw polysymmetric, anthers basifixed

References: Stevens PF (2019) APweb – www.mobot.org/MOBOT/research/APweb p
APG IV (2016); Judd W et al. (2016); Simpson M (2010); Soltis DE et al. (2005/2011/2014); Watson/Dallwitz (2018) a woody (except Apiaceae)
Apiaceae Griseliniaceae Pennantiaceae
© The Authors, 2019/2 (CC-BY)

A piales
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, n
7/494/5489

G inferior infl mostly umbel; drupe or schizocarp

Ray F. Evert, Mohamed Fennane, Marc Gottschling, Zigmantas Gudžinskas, Akitoshi Iwamoto, Chen-Kun Jiang, Anna Kagiampaki, Kent Kainulainen, Elizabeth A. Kellogg, Sangtae Kim,
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 (Apiaceae: mericarp/carpophore) Araliaceae Myodocarpaceae Pittosporaceae
fr usually
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 l
Paracryphiales
1–few-seeded
woody; infl racemose, flw 4-merous
i filaments stout; capsule septicidal Paracryphiaceae
1/3/36

Angiosperm Tracheophyte Bryophyte d lvs opposite, often basally connate

Phylogeny Phylogeny Phylogeny s buds with scales
Adoxaceae
flw often monosymmetric
Dipsacales
2/46/1090

Poster Poster Poster K persistent in fruit; secoiridoids Caprifoliaceae (incl. Dipsacaceae Diervillaceae Linnaeaceae Morinaceae Valerianaceae)