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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

2/10/100
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

4/17/4090
single adaxial prophyll; swollen nodes
y s
Calycanthaceae Hernandiaceae Monimiaceae
Laurales
woody; lvs opposite; flw with hypanthium, staminodes frequent
p

7/91/2858
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

6/154/2829
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
mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules

14/166/4560
woody; vessels absent
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
flw strobilar, perfect, parts free Monocots Ca oxalate
fr a follicle
East Asia Petrosaviales Petrosaviaceae

1/2/4
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

5/21/1037
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
flw tepals sometimes spotted, nectaries on tepals, anthers extrorse
Liliales

10/67/1558
siphonogamy lvs parallel-veined, entire
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

Arecales
lvs often palmately or pinnately pseudocompound, reduplicate-plicate

15/997/18875 2/192/2585
stem-borne roots numerous
intense primary growth, large apical meristem, infl often with spathe; alkaloids Arecaceae Dasypogonaceae
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 A often strongly modified/reduced, G inferior

8/92/2185
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

1/1/2
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
Gunnerales Gunneraceae Myrothamnaceae

2/2/50
ellagic acid
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
7/60/1815

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 Cannabaceae Moraceae Ulmaceae
9/261/7725

N fix
s s carpels with 1 ovule, stigma dry; dihydroflavonols
Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae)
embryo large
endosperm scanty lvs mostly alternate
flw often unisexual, G mostly inferior
Cucurbitales Apodanthaceae Begoniaceae Corynocarpaceae Datiscaceae
7/129/2320

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 Ticodendraceae
7/33/1055

ectomycorrhiza; tannins, dihydroflavonols Casuarinaceae Juglandaceae Nothofagaceae


stems sometimes jointed at nodes; lvs with glandular teeth

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


2(5)/17/836

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)
9/380/11731

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
9/471/6700

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 Cistaceae Cytinaceae Muntingiaceae Sarcolaenaceae


sperm
flw K often valvate, petals contorted, A often ∞; mucilage
10/338/6000

scanty
Liverworts cyclopropenoid fatty acids, flavones
Dipterocarpaceae Neuradaceae Sphaerosepalaceae Thymelaeaceae
Mosses
woody or herbs; flw often 4-merous Bataceae Caricaceae Limnanthaceae Salvadoraceae
Brassicales
18/398/4765

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
37/749/11600

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 Polemoniaceae Sarraceniaceae
s lvs teeth often theoid; nodes unilacunar

Ericales
22/346/11545

embryo flw 5-merous, pentacyclic Balsaminaceae Fouquieriaceae Primulaceae Styracaceae


Asterids Campanulids t long nonhydrolyzable tannins, ellagic acid, triterpenoids, hydroquinones
Clethraceae Lecythidaceae Roridulaceae Theaceae
e
r
tropical trees; lvs exstipulate/entire; nodes often unilacunar Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae
pedicels articulated; style short; embryo long
i endosperm copious
Icacinales Oncothecaceae Icacinaceae
2/24/202

d indole and quinoline alkaloids (camptothecin)

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; Africa to India; lvs opposite, glandular-pubescent
s
Vahliales
mostly sympetalous flw < 5 mm across, K valvate, G(3) inf, nectary
Peter F. Stevens, Ph.D.
C enclosing fr septicidal capsule, K/C/A persistent; seeds minute; iridoids Vahliaceae
1/1/8

A and G in bud
Missouri Botanical Garden (MoBot) G(2) lvs opposite; interpetiolar stipules or nodal line/ridge
C Apocynaceae (incl. Asclepiadaceae) Loganiaceae
Gentianales
style single, long colleters
5/1118/19840

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 iridoids + A
lvs opposite; nodes 1:1; flw mostly monosymmetric
Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae
A often 2(+2); gland-headed hairs with radially arranged cells
Lamiales
24/1059/23810

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 (2018) Angiosperm Phylogeny Poster – Flowering Plant Systematics
woody; lvs serrate
• hypothetical tree based on molecular phylogenetic data (Dec. 2016)
• 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
• numbers set in gray next to orders refer to families, genera, and species C mostly woody; infl racemose, C free
Escalloniales Escalloniaceae
1/9/130

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 (2018) 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
A piales
lvs often divided; nodes usu. multilacunar
© The Authors, 2017 (CC-BY)

Acknowledgements to our contributors, translators, and consultants: Julien B. Bachelier, Przemysław Baranow, Zoltán Barina, Fernanda Antunes Carvalho, Christoph Dobeš, Ray F. Evert,
n
7/494/5489

G inferior infl mostly umbel; drupe or schizocarp


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, Peter H. Raven, u (Apiaceae: mericarp/carpophore) Araliaceae Myodocarpaceae Pittosporaceae
fr usually
Yasaman Salmaki, Federico Selvi, Paramjit Singh, Douglas E. Soltis, Pramote Triboun, Zoya M. Tsymbalyuk, Magsar Urgamal, Maximilian Weigend, Michael Wink, Shahin Zarre
l 1–few-seeded

i
woody; infl racemose, flw 4-merous
filaments stout; capsule septicidal Paracryphiales 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)