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systematics, 2017
The "Angiosperm Phylogeny Poster Flowering Plant Systematics" is an educational tool presenting an
overview of the evolutionary relationships among flowering plants according to APG IV and APweb as of
2016. The phylogenetic tree depicts the 64 orders and the majority of the 416 families, listing the main
apomorphies and plesiomorphies as well as diagnostic and nondiagnostic anatomical, morphological, and
phytochemical features for orders and higher levels within the tree. The intuitive color-coding facilitates
memorization and teaching.
Translations of the poster by internationally renown botanists are now available in 18 languages.
Hyperlinks to APweb (Peter F. Stevens, Missouri Botanical Garden) are provided for the orders and higher
ranks.
This is one in a series of three educational posters on the phylogeny of land plants: Poster 1:
"Angiosperms", Poster 2: "Tracheophytes: Lycophytes, Ferns, Gymnosperms", Poster 3: "Bryophytes:
Liverworts, Mosses, Hornworts".
PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2320v3 | CC BY 4.0 Open Access | rec: 9 Jan 2017, publ: 9 Jan 2017
Angiosperm Phylogeny
Flowering Plant Systematics
A
woody, vessels lacking; dioecious; flw T58, A, G58, 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 T412, A1, embryo sac 4-nucleate
r
seeds operculate with perisperm but endosperm reduced or small; mucilage; alkaloids (no benzylisoquinolines) Cabombaceae Hydatellaceae Nymphaeaceae
n
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
tiglic acid, aromatic terpenoids
A e
chlorAnthAles
lvs opposite, interpetiolar stipules; nodes swollen
A flw small T03, A15, G1, 1 apical ovule/carpel Chloranthaceae
1/4/75
e n
A G woody; foliar sclereids, K and C distinct
cAnellAles
sesquiterpenes
2/10/100
aromatic terpenoids Canellaceae Winteraceae
r i
tension wood +
(pellucid dots)
nodes trilacunar
l o herbaceous; lvs two-ranked, leaf base sheathing
piperAles
4/17/4090
single adaxial prophyll; swollen nodes Aristolochiaceae (incl. Hydnoraceae) Piperaceae Saururaceae
y s
woody; lvs opposite; flw with hypanthium, staminodes frequent Calycanthaceae Hernandiaceae Monimiaceae
lAurAles
7/91/2858
p often valvate anthers; carpels with 1 ovule; embryo large
Gomortegaceae Lauraceae Siparunaceae
e mAGnoliiDs
benzylisoquinoline alkaloids
mAGnoliAles
woody; pith septate; lvs two-ranked; ovules with obturator Annonaceae Eupomatiaceae Magnoliaceae
6/154/2829
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
s ovules atropous, seeds with epidermal perisperm and copious endosperm; ethereal oils Acoraceae
1/1/2-4
Alismataceae Butomaceae Posidoniaceae Scheuchzeriaceae
AlismAtAles
14/166/4560
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
1/2/4
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
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
5/36/1345
G apocarpous (style short in most) sieve tube plastids with infl sometimes spathe + spadix Cyclanthaceae Pandanaceae Triuridaceae Velloziaceae
compitum (if present) extragynoecial protein crystals
nectaries absent often geophytes (bulbs, tubers, rhizomes); leaf bases often not sheathing
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
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
mostly herbaceous; epidermis siliceous; mostly mycorrhiza absent Bromeliaceae Eriocaulaceae Poaceae Restionaceae Xyridaceae
lvs grassy; flw often anemophilous, minute, chaffy, without nectaries poAles 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 910*, 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
recepta-
e 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
2/2/50
ellagic acid
c
absent
1/10/300
t seeds often with aril; fr usually follicles
benzylisoquinolines
s lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent
Altingiaceae Cynomoriaceae Haloragaceae Peridiscaceae
sAxifrAGAles
15/112/2500
fr mostly dry, dehiscent
pollen tricolpate
protandry common
myricetin, flavonols 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
resinous, lignans/neolignans, harman alkaloids
2/24/345
seeds often arillate (red-orange) or winged Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae) Lepidobotryaceae
s lvs often compound, pulvini (sleep movement)
u Brunelliaceae Connaraceae Elaeocarpaceae
COM clade flw A5 or multiple, branched style common
oxAliDAles Oxalidaceae
7/60/1815
fAbAles
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)
Ticodendraceae
ectomycorrhiza; tannins, dihydroflavonols Casuarinaceae Juglandaceae Nothofagaceae
stems sometimes jointed at nodes; lvs with glandular teeth
GerAniAles
2(5)/17/836
flw A obdiplostemonous, nectary outside A; fr capsule Geraniaceae Francoaceae (incl. Ledocarpaceae, Melianthaceae, Vivianiaceae)
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
pollen tricolporate
e endosperm scanty, scaly bark, flavonols, myricetin Melastomataceae (incl. Memecylaceae) Onagraceae Vochysiaceae
G connate n woody
style + t stylodia free
crossosomAtAles Crossosomataceae Stachyuraceae
Strasburgeriaceae
7/12/66
ferns
(incl. horsetAils) woody; (semi-)parasites; mycorrhiza absent; lvs margins entire
flw A epipetalous; perianth often simple, valvate, persisting Balanophoraceae Misodendraceae Opiliaceae Schoepfiaceae
sAntAlAles
13/151/1992
cAryophyllAles
conifers
plAnts Gymnosperms
AnA GrADe s pollen colpate, surface spiny
betalains or anthocyanins (latter, e.g., in Caryophyllaceae)
(incl. Chenopodiaceae)
Basellaceae
Droseraceae
Drosophyllaceae
Nyctaginaceae
Phytolaccaceae
Simmondsiaceae
Talinaceae
mAGnoliiDs u mostly woody; lvs mostly undivided, hydathode teeth Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae
p flw often 4-merous, K much smaller than C, persisting
monocots Cornaceae Grubbiaceae Loasaceae
e intrastaminal disk, G inferior; fr drupaceous
cornAles
6/51/590
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
t woody; dioecious
Theodor C. H. Cole, Dipl. Biol. e
Hartmut H. Hilger, Prof. Dr. r l
flw small, C valvate, G unilocular; fruit indehiscent
iridoids (aucubin), gutta GArryAles Eucommiaceae Garryaceae (incl. Aucubaceae)
2/3/18
Dahlem Centre of Plant Sciences (DCPS) i A lvs spiral, simple; nodes unilacunar
m Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae)
Institute of Biology Botany D solAnAles
5/165/4125
A and G in bud
G(2)
Peter F. Stevens, Ph.D. style single, long C lvs opposite, colleters Apocynaceae (incl. Asclepiadaceae) Loganiaceae
GentiAnAles
5/1118/19840
and infl A often 2(+2); gland-headed hairs with radially arranged cells
lAmiAles
epipetalous
cymose 6-oxygenated flavones, Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae
University of MissouriSt. Louis fr a drupe oligosaccharides cornoside, verbascoside (acetoside) Byblidaceae Martyniaceae Pedaliaceae Stilbaceae
seed single
Department of Biology ellagic acid lvs roughly hairy; nodes unilacunar Gesneriaceae Oleaceae Phrymaceae Verbenaceae
St. Louis, MO 63121-4499, USA lacking
infl scorpioid; mostly 4 ovules
6-11/150/3095
COLE TCH, HILGER HH, STEVENS PF (2017) Angiosperm Phylogeny Poster Flowering Plant Systematics fr usually drupe with broad stigma
hypothetical tree based on molecular phylogenetic data (Dec. 2016) early
phylogeny, classification, and features chiefly follow APweb and APG
sympetaly
flw small
herbs or shrubs; flw often monosymmetric Asteraceae Goodeniaceae Pentaphragmataceae
plunger sec. pollination device (long style)
AsterAles
11/1743/26870
64 orders and around 420 families currently recognized by APG IV (some minor families excluded here)
contrary to APG, but in compliance with APweb and other seminal sources, we here recognize several families within Boraginales
embryo nodes trilacunar Calyceraceae Menyanthaceae Rousseaceae
short
branch lengths deliberate, not expressing actual time scale inulin, sesquiterpenes, secoiridoids Campanulaceae (incl. Lobeliaceae) Stylidiaceae
the characters listed do not necessarily apply to all members of the according clade
position of various characters on the tree uncertain c mostly woody; infl racemose, C free
escAlloniAles Escalloniaceae
1/9/130
for family characteristics see: FGVP, Kubitzki K, ed. (1990 ff) anthers basifixed, nectary disc
orders (and higher ranks) are linked to the according pages on APweb A
numbers set in gray next to orders refer to families, genera, and species m woody; evergreen; nodes 1:1
bruniAles Bruniaceae Columelliaceae (incl. Desfontainia)
2/14/79
l 1few-seeded
pArAcryphiAles
Akitoshi Iwamoto, Chen-Kun Jiang, Anna Kagiampaki, Elizabeth A. Kellogg, Sangtae Kim, Asl Doru-Koca, Nikos Krigas, Diego Medan, Sergei L. Mosyakin, Sofi Mursidawati, Anastasiya V. Odintsova, woody; infl racemose, flw 4-merous
Richard G. Olmstead, Peter H. Raven, Yasaman Salmaki, Federico Selvi, Douglas E. Soltis, Trn Hu ng, Pramote Triboun, Zoya M. Tsymbalyuk, Maximilian Weigend, Michael Wink, Shahin Zarre i filaments stout; capsule septicidal Paracryphiaceae
1/3/36
Poster Poster Poster K persistent in fruit; secoiridoids (incl. Dipsacaceae Diervillaceae Linnaeaceae Morinaceae Valerianaceae)
PeerJ Preprints | https://doi.org/10.7287/peerj.preprints.2320v3 | CC BY 4.0 Open Access | rec: 9 Jan 2017, publ: 9 Jan 2017