Angiosperm Phylogeny

Flowering Plant Systematics

A g
woody, vessels lacking; dioecious; flw T5–8, A∞, G5–8; 1 ovule/carpel; embryo sac 9-nucleate; 1 species (New Caledonia) A mborellales Amborellaceae
N
I
r
a
aquatic, herbaceous; cambium absent; aerenchyma; flw T4–12, A3–∞; embryo sac 4-nucleate
seeds operculate, perisperm; mucilage; alkaloids (no benzylisoquinolines) N ymphaeales Cabombaceae Hydatellaceae Nymphaeaceae

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

T d
e
tiglic acid, aromatic terpenoids Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae
A
Chloranthales
lvs opposite, interpetiolar stipules; nodes swollen
e flw small T0–3, A1–5, G1, 1 apical ovule/carpel Chloranthaceae
a
Canellales
woody; pollen uniporate
sesquiterpenes

r aromatic terpenoids Canellaceae Winteraceae

l
(pellucid dots)

nodes trilacunar
Piperales
± herbaceous; lvs two-ranked, leaf base sheathing
y single adaxial prophyll; swollen nodes Aristolochiaceae Hydnoraceae Piperaceae Saururaceae
Calycanthaceae Hernandiaceae Monimiaceae
Laurales
woody; lvs opposite; flw with hypanthium, staminodes frequent
a often valvate anthers; carpels with 1 ovule; embryo large
Gomortegaceae Lauraceae Siparunaceae
n Magnoliids
benzylisoquinoline alkaloids;

Magnoliales Annonaceae Eupomatiaceae Magnoliaceae

woody; pith septate; lvs two-ranked; ovules with obturator
g features as in endosperm ruminate
Degeneriaceae Himantandraceae Myristicaceae
embryo sac 8-nucleate

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

o ovules atropous, with epidermal perisperm and copious endosperm; idioblasts with ethereal oils Acoraceae
s Alismataceae (incl. Limnocharitaceae) Juncaginaceae Ruppiaceae
Alismatales
mostly herbs and aquatics; rhizomatous; hydrophilous; intravaginal squamules
Aponogetonaceae Butomaceae Posidoniaceae Scheuchzeriaceae
p flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green
Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae
e
Petrosaviales
stem with ring of bundles
r Monocots Ca oxalate
raphides
fr a follicle; East Asia Petrosaviaceae
m
Dioscoreales
endosperm
atactostele (scattered bundles) often twining vines; lvs often reticulate
nuclear Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae
s no secondary thickening
mostly herbaceous
helobial
nectaries
ovary often inferior, style short, branched; steroid sapogenins/alkaloids

some woody (with terminally tufted leaves)

Pandanales
when present
pollen monosulcate
woody; vessels absent sieve tube plastids with
septal
flw spadix Cyclanthaceae Pandanaceae Velloziaceae
eustele; nodes protein crystals
sieve tube plastids often geophytes (bulbs, tubers, rhizomes)
sympodial branching
Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae
with starch grains
lvs simple, persistent, entire
flw parts free, strobilar,
vascular bundles in stem scattered
lvs parallel-veined, entire
no glandular teeth
flw tepals sometimes spotted, nectaries at tepals; many seeds, seeds coat (testa) cellular
phytomelan lacking; fructans in stems, chelidonic acid, steroid saponins Liliales Colchicaceae Liliaceae Petermanniaceae Smilacaceae
perfect, P parts varying, flw pentacyclic, P 3-merous, A opp. P often geophytes; anomalous sec. growth Amaryllidaceae ( incl. Agapanthaceae , Alliaceae) Hypoxidaceae Iridaceae
Asparagales
often in threes, filaments narrow capsule or berry
weakly differentiated,
stamen with broad filaments
anthers broadly attached,
septal nectary; single cotyledon
seed coat obliterated or with phytomelan Asparagaceae ( incl. Agavaceae, Hyacinthaceae, Ruscaceae) Lanariaceae Orchidaceae
pollen monosulcate adventitious roots Tecophilaeaceae Xanthorrhoeaceae ( incl. Asphodelaceae, Hemerocallidaceae)
G apocarpous (style short in most) monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate

Arecales
nectaries absent intense primary growth, large apical meristem
embryo very small infl with spathe; alkaloids Arecaceae
aromatic terpenoids

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 often irregular; few fertile stamens
Commelinids infl thyrsus of scorpioid cymes
phenylphenalenones Commelinales Commelinaceae Haemodoraceae Pontederiaceae
UV-fluorescing cell walls rhizomatous, large-leafed herbs; pseudostem: central infl
(ferulic/coumaric acids) flw irregular/monosymmetric, septal nectaries
silicic acid in leaves
cuticular waxes often in rodlets
aggregated into scallops
G inferior, A often strongly modified/reduced
seeds arillate; silicic acid Zingiberales Cannaceae
Costaceae
Heliconiaceae
Lowiaceae
Marantaceae
Musaceae
Strelitziaceae
Zingiberaceae

Ceratophyllales
aquatic, herbaceous; lvs whorled, no pellucid dots; vessels lacking; monoecious
flw T9 10, A∞, G1, 1 apical ovule/carpel, pollen inaperturate, pollen tube branched; hydrophilous
– Ceratophyllaceae
Berberidaceae Eupteleaceae Menispermaceae
Ranunculales
lvs often divided; flw whorled, P single or multiple whorls
G apocarpous/paracarpous, superior; berberines Ranunculaceae
Circaeasteraceae Lardizabalaceae Papaveraceae

Sabiales
woody; flw 4–5-merous, nectary: disk, A epipetalous, G syncarpous
pollen colporate; endosperm helobial; triterpenoids Sabiaceae

Proteales
mostly woody; flw tepals often 4-merous
A epitepalous, connectives sometimes with apical appendage Nelumbonaceae Platanaceae Proteaceae
E axial woody; vessels lacking; idioblasts in cortex; flw tepals missing, A∞

u
nectary
common
G>5 laterally connate with abaxial nectaries; fr aggregate of follicles Trochodendrales Trochodendraceae
d
Buxales
mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic
Buxaceae Didymelaceae
benzylisoquinolines absent

i flw tepals ± uniform or missing; pregnan pseudoalkaloids

c unisexual, dioecious; lvs toothed, sec. veins palmate

o
t
flw tepals small to lacking
ellagic acid Gunnerales Gunneraceae Myrothamnaceae
mostly woody; lvs veins proceed to apex & teeth; flw K5–∞, persisting
s
pollen tricolpate
mostly A∞, G mostly slightly connate
seeds often with aril; fr often both ventrally/dorsally dehiscent Dilleniales Dilleniaceae
flw K/C/P opp A lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent
Altingiaceae Daphniphyllaceae Hamamelidaceae
Saxifragales
filaments rather narrow fr mostly dry, dehiscent
anthers basifixed
nodes trilacunar
myricetin, flavonols Cercidiphyllaceae Grossulariaceae Paeoniaceae
stomata anomocytic often tendrillar vines; lvs often divided and with glandular teeth Crassulaceae Haloragaceae Saxifragaceae
ethereal oils absent

Vitales
A epipetalous, 2 ovules per carpel; raphides, pearl glands
stipules
berries Vitaceae
nodes cork origin deep-seated

Zygophyllales
3:3
endosperm lacking
resinous, lignans/neolignans, harman alkaloids Krameriaceae Zygophyllaceae
ellagic acid tannins

R flw small, G often 3-merous, nectary: intrastaminal disk

o
s
seeds often with aril (red-orange)
infl cymose Celastrales Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae) Lepidobotryaceae
i lvs often compound, pulvini (sleep movement)
Brunelliaceae Connaraceae Elaeocarpaceae
d
s
flw A5 or multiple, branched style common
mucilage cells; oxalates Oxalidales Cephalotaceae Cunoniaceae Huaceae
Oxalidaceae

F Achariaceae Euphorbiaceae Rafflesiaceae Ochnaceae Podostemaceae

a lvs margins toothed
flw G often tricarpellate Malpighiales Chrysobalanaceae
Clusiaceae
Hypericaceae
Linaceae
Passifloraceae
Phyllanthaceae


Rhizophoraceae
Salicaceae
b
i flw often “papilionaceous”: wing, standard, keel, mostly G1 Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae

Fabales
mostly A10; fr a pod; symbiosis with root nodule bacteria
d diverse alkaloids, NP amino acids, lectins (in Fabaceae) Fabaceae Polygalaceae Quillajaceae Surianaceae
c E s
u lvs mostly simple with stipules Barbeyaceae Elaeagnaceae Rosaceae
o
r d N fix flw K valvate (and hypanthium) persisting
carpels with 1 ovule, stigma dry; dihydroflavonols Rosales Cannabaceae Moraceae Ulmaceae
e i embryo large Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae)
c endosperm scanty lvs mostly alternate

Cucurbitales Anisophyllaceae Coriariaceae Cucurbitaceae

flw often unisexual, G mostly inferior
o parietal placentation; cucurbitacins Tetramelaceae
t lvs undivided; flw small, unisexual
Begoniaceae Corynocarpaceae Datiscaceae
s anemophilous, thus T reduced or lacking, G mostly inferior
Betulaceae Fagaceae Myricaceae Rhoipteleaceae
flw pentacyclic
parts alternating
infl spikes or catkins; fr 1-seeded, mostly nuts
ectomycorrhiza; tannins, dihydroflavonols Fagales Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae
lvs with glandular teeth

Geraniales
G connate
K + C (free)
flw A obdiplostemonous, nectary on filament
stems jointed at nodes; ethereal oils, ellagic acid
Francoaceae Geraniaceae Ledocarpaceae Melianthaceae
(A polyandrous)
pollen tricolporate lvs opposite, colleters (glandular hair on adaxial surface of petiole base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae)
trihydroxyl-flavonoids stipules small (if any); cork deep seated
Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae)
flw K valvate, persisting, A incurved in bud, ovary inferior; ovules many
endosperm scanty; scaly bark; flavonols, myricetin Myrtales Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae
woody
K + C, stylodia free
hypanthium, nectary disk Crossosomatales Crossosomataceae
Geissolomataceae
Stachyuraceae
Staphyleaceae
Strasburgeriaceae

M
trees; lvs spiral; extrafloral nectaries

a
staminate flw: A = and opposite C
bark bitter, anthraquinones Picramniales Picramniaceae
mostly woody (silica/silicified)
l
v
lvs alternate, odd-pinnately compound
flw often imperfect, intrastaminal disk; ethereal oils Sapindales Anacardiaceae
Burseraceae
Meliaceae
Nitrariaceae
Rutaceae
Sapindaceae
Simaroubaceae

i vessel elements: scalariform perforations; mucilage cells
d
s
embryo
large
lvs margins toothed, stipules cauline
flw small, A = and opposite K, ovules 1-2/carpel Huerteales Dipentodontaceae Gerrardinaceae Petenaeaceae Tapisciaceae
endo- bark fibrous; hairs often stellate Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
sperm flw K often valvate
scanty petals contorted, A often ∞; mucilage
cyclopropenoid fatty acids, flavones Malvales Cistaceae
Dipterocarpaceae
Cytinaceae
Neuradaceae
Muntingiaceae
Sphaerosepalaceae
Sarcolaenaceae
Thymelaeaceae
Liverworts lvs alternate; flw often 4-merous
often clawed petals, A often many, 2×K Bataceae Caricaceae Limnanthaceae Salvadoraceae
Mosses hypogynous (often gynophore); infl racemose
myrosin cells, glucosinolates Brassicales Brassicaceae Cleomaceae Moringaceae Tovariaceae
Hornworts woody; (semi-)parasites; without mycorrhiza; lvs margins entire Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae
flw A epipetalous; perianth often simple, valvate, persisting
Lycophytes

Ferns
free-central pendulous placentation; fr one-seeded
polyacetylenes, triterpene sapogenins, silicic acid Santalales Balanophoraceae
Loranthaceae
Misodendraceae
Olacaceae
Opiliaceae
Santalaceae
Schoepfiaceae
Viscaceae
(incl. horsetails) woody; lvs stomata cyclocytic
Palmferns
Ginkgo
petiole bundles annular; fr fleshy
calcium oxalate as crystals Berberidopsidales Aextoxicaceae
Aizoaceae
Berberidopsidaceae
Caryophyllaceae Molluginaceae Polygonaceae
Ephedra mostly herbaceous; without mycorrhiza
Welwitschia G often unilocular with central placentation, Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae
Seed Gymnosperms
Caryophyllales [incl. Chenopodiaceae] Droseraceae Nyctaginaceae Simmondsiaceae
Gnetum pollen colpate, surface spiny
Conifers
Plants betalains or anthocyanins (latter, e.g., in Caryophyllaceae) Basellaceae Drosophyllaceae Phytolaccaceae Talinaceae
ANITA grade Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae
mostly woody; lvs mostly undivided, hydathode teeth
Magnoliids flw often 4-merous, K much smaller than C, persisting
Cornaceae Grubbiaceae Loasaceae
Angiosperms Monocots
intrastaminal disk, G inferior; fr drupaceous
diverse iridoids Cornales Curtisiaceae Hydrangeaceae Nyssaceae
Fabids
lvs teeth, theoid; nodes unilacunar Actinidiaceae Ericaceae Polemoniaceae Sarraceniaceae
Balsaminaceae Fouquieriaceae Primulaceae Styracaceae
Ericales
Malvids flw 5-merous, pentacyclic
Rosids nonhydrolyzable tannins, ellagic acid, hydroquinones
Lamiids
Clethraceae Lecythidaceae Roridulaceae Theaceae
tropical trees; lvs exstipulate/entire; nodes often unilacunar
pedicels articulated; style short; embryo long
Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae
Campanulids
Icacinales
Asterids fr 1-seeded (drupe); endosperm copious
indole and quinoline alkaloids (camptothecin) Oncothecaceae Icacinaceae
tropical trees; few common features; nodes usu. trilacunar

A late
sympetaly
Metteniusa: NE Andes, cloud montane forest; lvs alternate; flw bisexual, fragrant; anthers sagittate,
pollen sacs moniliform, dehisce longitudinally; infl axillary cyme; G(5) unilocular; fr 1-seeded (drupe) Metteniusales Metteniusaceae
s woody; dioecious
Theodor C. H. Cole, Dipl. Biol.
Institute of Pharmacy and Molecular Biotechnology
t
e L
flw small, C valvate, G unilocular; fruit indehiscent
iridoids (aucubin), gutta Garryales Eucommiaceae Garryaceae (incl. Aucubaceae)
Heidelberg University
r a lvs opposite, colleters
Apocynaceae (incl. Asclepiadaceae) Loganiaceae
Im Neuenheimer Feld 364
D-69120 Heidelberg, Germany
i
d
m
ii
flw corolla convolute in bud
indole alkaloids; iridoids Gentianales Gentianaceae Gelsemiaceae Rubiaceae
s d lvs spiral, simple; nodes unilacunar
Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae)
mostly sympetalous
A=C,
epipetalous
s flw petals plicate; K persisting
diverse alkaloids, no iridoids Solanales Hydroleaceae Montiniaceae Sphenocleaceae
nectary gynoecial
ovules unitegmic
polyandry
rare,
lvs opposite; nodes 1:1; flw mostly monosym. Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae
A often 2(+2); ethereal oils in gland-headed hairs
endosperm cellular G(2) Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae
iridoids common
infl
cymose
6-oxygenated flavones, rosmarinic acid,
oligosaccharides: cornoside, verbascoside (acetoside) Lamiales Byblidaceae Martyniaceae Pedaliaceae Stilbaceae
ellagic acid lvs roughly hairy; nodes unilacunar Gesneriaceae Hydrostachyaceae Oleaceae Phrymaceae Verbenaceae
lacking infl scorpioid; mostly 4 ovules

Prof. Dr. Hartmut H. Hilger

isokestose, higher inulins,
pyrrolizidine alkaloids Boraginales Boraginaceae Codonaceae Coldeniaceae Cordiaceae Ehretiaceae (+ Lennoaceae)
Heliotropiaceae Hydrophyllaceae Namaceae Wellstediaceae
Dahlem Centre of Plant Sciences (DCPS) woody; lvs serrate
Institute of Biology – Plant Morphology and Systematics
Freie Universität Berlin early
sympetaly
flw 1-2 ovules/carpel, C± free, K slightly connate
fr drupe with broad stigma Aquifoliales Aquifoliaceae Cardiopteridaceae Stemonuraceae
Altensteinstr. 6, D-14195 Berlin, Germany flw small flw often monosymmetric Asteraceae Goodeniaceae Pentaphragmataceae
plunger pollination (long style)
Asterales
________________________________________________________________________________________ embryo
short nodes trilacunar Calyceraceae Menyanthaceae Rousseaceae
- hypothetical tree based on molecular phylogenetic data (Oct. 2015) inulin, sesquiterpenes, secoiridoids Campanulaceae (incl. Lobeliaceae) Stylidiaceae
- phylogeny, classification, and features chiefly follow APweb and APG
- approx. 450 families currently recognized (some minor families excluded here)
- branch lengths deliberate, not expressing actual time scale C mostly woody; infl racemose, C free
anthers basifixed, nectary disc Escalloniales Escalloniaceae
- if a character is marked as being a potential synapomorphy at a node/for a clade; a
m Bruniales
this does not mean that all members of that clade possess that character woody, evergreen; nodes 1:1
- position of various characters on the tree unclear flw polysymmetric, anthers basifixed Bruniaceae Columelliaceae (incl. Desfontainia)
- References: Stevens PF (2015) APweb – www.mobot.org/MOBOT/research/APweb p woody (except Apiaceae)
APG III (2009); Judd W et al. (2007); Simpson M (2010); Soltis DE et al. (2005/2011/2014); Watson/Dallwitz (2015) delta-intkey.com/ a lvs often divided; nodes usu. multilacunar
Apiaceae Griseliniaceae Pennantiaceae
- for family characteristics see: Kubitzki K, ed. (1990 ff) n
u
G inferior
fr usually
infl mostly umbel; drupe or schizocarp
(Apiaceae: mericarp/carpophore) A piales Araliaceae Myodocarpaceae Pittosporaceae
Special thanks for ongoing consultation since inception of this poster to Peter F. Stevens (Missouri Botanical Garden)
l 1–few-seeded

Paracryphiales
Acknowledgements to Christoph Dobeš, Ray F. Evert, Marc Gottschling, Richard G. Olmstead, Peter H. Raven, Douglas E. Soltis, Maximilian Weigend, Michael Wink woody; infl racemose, flw 4-merous,
i filaments stout, capsule septicidal Paracryphiaceae
Angiosperm Tracheophte Bryophte d lvs opposite, often basally connate
Phylogeny Phylogeny Phylogeny s nodes 3:3; buds with scales
Adoxaceae Diervillaceae Linnaeaceae
Poster Poster Poster flw often monosymmetric
K persistent in fruit; secoiridoids Dipsacales Caprifoliaceae Dipsacaceae Morinaceae
Valerianaceae