Angiosperm phylogeny poster (APP) – Flowering plant systematics, 2019

Abstract

NEW UPDATE! The ANGIOSPERM PHYLOGENY POSTER (APP) – Flowering Plant Systematics is an educational tool presenting an overview of the evolutionary relationships among flowering plants according to APG IV, APweb, and PPA (Li HT et al. Nature Plants 2019). 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 along with the number of families, genera, and species for each order. The intuitive color-coding facilitates memorization and teaching.
Translations of the poster by internationally renowned botanists are now several 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".

Full text

AsterAles
Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
Cistaceae Cytinaceae Muntingiaceae
Sarcolaenaceae
Dipterocarpaceae Neuradaceae Sphaerosepalaceae
Thymelaeaceae
bark brous; hairs often stellate
w K often valvate, petals contorted, A often ∞; mucilage
cyclopropenoid fatty acids, avones
herbs or shrubs; nodes trilacunar
w often monosymmetric
plunger sec. pollination device (long style)
inulin, sesquiterpenes, secoiridoids
Asteraceae Goodeniaceae
Pentaphragmataceae
Calyceraceae Menyanthaceae
Rousseaceae
Campanulaceae
(
incl.
Lobeliaceae
)
Stylidiaceae
endo-
sperm
scanty
DipsAcAles
Adoxaceae
Caprifoliaceae (incl. Dipsacaceae Diervillaceae Linnaeaceae Morinaceae Valerianaceae)
AmborellAles
ApiAles
Apiaceae Griseliniaceae
Pennantiaceae
Araliaceae
Myodocarpaceae
Pittosporaceae
AquifoliAles
Aquifoliaceae Cardiopteridaceae
Stemonuraceae
GArryAles
Eucommiaceae Garryaceae (incl. Aucubaceae)
ericAles
cornAles
Cornaceae Grubbiaceae
Loasaceae
Curtisiaceae Hydrangeaceae Hydrostachyaceae
Nyssaceae
mAlvAles
brAssicAles
cucurbitAles
rosAles
fAbAles
mAlpiGhiAles
celAstrAles
myrtAles
GerAniAles
crossosomAtAles
cAnellAles
piperAles
lAurAles
mAGnoliAles
vitAles
sAxifrAGAles
GunnerAles
Bataceae Caricaceae Limnanthaceae
Salvadoraceae
Brassicaceae Cleomaceae Moringaceae
Tovariaceae
Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae
fAGAles
Betulaceae Fagaceae Myricaceae
Casuarinaceae Juglandaceae Nothofagaceae
Ticodendraceae
Apodanthaceae Begoniaceae Corynocarpaceae Datiscaceae
Anisophyllaceae
Coriariaceae Cucurbitaceae
Tetramelaceae
nymphAeAles
A
ustrobAileyAles
ZyGophyllAles
DilleniAles
Fabaceae Polygalaceae Quillajaceae
Surianaceae
Barbeyaceae Elaeagnaceae Rosaceae
Cannabaceae Moraceae Ulmaceae
Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae)
Celastraceae
(incl.
Hippocrateaceae, Brexiaceae, Parnassiaceae
)
Lepidobotryaceae
Achariaceae Euphorbiaceae Rafesiaceae Ochnaceae Podostemaceae
Chrysobalanaceae Hypericaceae Passioraceae Rhizophoraceae
Clusiaceae Linaceae Phyllanthaceae Salicaceae
Erythroxylaceae Malpighiaceae Picrodendraceae
Violaceae
Krameriaceae Zygophyllaceae
Combretaceae
Myrtaceae Penaeaceae (incl. Oliniaceae)
Lythraceae (incl. Punicaceae, Sonneratiaceae, Trapaceae)
Melastomataceae (incl. Memecylaceae) Onagraceae
Vochysiaceae
Geraniaceae
Francoaceae (incl.
Ledocarpaceae, Melianthaceae, Vivianiaceae
)
Crossosomataceae
Stachyuraceae
Strasburgeriaceae
Geissolomataceae
Staphyleaceae
Vitaceae
m
AGnoliiDs
Dilleniaceae
Gunneraceae Myrothamnaceae
Amborellaceae
Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae
Cabombaceae Hydatellaceae Nymphaeaceae
Canellaceae Winteraceae
Annonaceae Eupomatiaceae Magnoliaceae
Degeneriaceae Himantandraceae Myristicaceae
Calycanthaceae Hernandiaceae
Monimiaceae
Gomortegaceae Lauraceae Siparunaceae
Aristolochiaceae (incl. Hydnoraceae) Piperaceae
Saururaceae
woody; vessels lacking
dioecious; w T5–8, A∞, G5–8, ovule 1/carpel, embryo sac 9-nucleate
1 species, New Caledonia
herbaceous, aquatic; cambium absent; aerenchyma; w T4–12, A1–∞, embryo sac 4-nucleate
seeds operculate, perisperm
mucilage; alkaloids (no benzylisoquinolines)
woody, vessels solitary
w T>10, A∞, G ca.9, embryo sac 4-nucleate
tiglic acid, aromatic terpenoids
woody; foliar sclereids
K and C distinct
aromatic terpenoids
± herbaceous; lvs two-ranked, leaf base sheathing
single adaxial prophyll
nodes swollen
woody; pith septate; lvs two-ranked
ovules with obturator
endosperm ruminate
woody; lvs opposite
w with hypanthium, staminodes frequent
anthers often valvate; ovule 1/carpel; embryo large
mostly herbaceous; without mycorrhiza
G often unilocular with free-central placentation
pollen colpate, surface spiny
betalains or anthocyanins (latter, e.g., in Caryophyllaceae)
lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent
fr mostly dry, dehiscent
myricetin, avonols
stems jointed at nodes (in some); lvs with glandular teeth
w A obdiplostemonous, nectary outside A; fr capsule
ethereal oils, ellagic acid
lvs opposite, colleters (glandular hair on adaxial surface of petiole base)
stipules small (if any), cork deep seated
w K valvate, persisting; A incurved in bud, ovary inferior, ovules many
endosperm scanty, scaly bark; avonols, myricetin
in cymose, w small
G often 3-merous, nectary often intrastaminal disk
seeds often arillate (red-orange) or winged
habits and habitats extremely diverse
lvs margins toothed; pedicels often articulated
w G3 (often)
w
often
“papilionaceous”: wing, standard, keel, C clawed, G1
(most)
, A10
(most)
fr pod
symbiosis with root nodule bacteria
diverse alkaloids, NP amino acids, lectins (in Fabaceae)
lvs mostly simple with stipules
w K valvate (and hypanthium) persisting
stigma dry, ovule 1/carpel
dihydroavonols
lvs mostly alternate
w often unisexual, G mostly inferior
parietal placentation
cucurbitacins
mostly trees; lvs mostly undivided; w small, unisexual
anemophilous, thus T reduced or lacking, G mostly inferior
in spikes or catkins; fr mostly nuts; seed 1
ectomycorrhiza; tannins, dihydroavonols
woody or herbs
in racemose; w often 4-merous, petals often clawed
myrosin cells, glucosinolates
mostly woody; lvs mostly undivided, hydathode teeth
w often 4-merous, K much smaller than C, persisting
intrastaminal disk, G inferior; fr drupaceous
diverse iridoids
mostly sympetalous
C enclosing
A and G in bud
G(2)
style single, long
nectary gynoecial
ovules unitegmic
endosperm cellular
iridoids +
lvs teeth often theoid; nodes unilacunar
w 5-merous, pentacyclic
nonhydrolyzable tannins, ellagic acid
triterpenoids, hydroquinones
late
sympetaly
woody; lvs serrate
w 1-2 ovules/carpel, C ± free, K slightly connate
fr usu. drupe with broad stigma
woody (except Apiaceae)
lvs often divided; nodes usu. multilacunar
in mostly umbel; fr drupe or schizocarp
(Apiaceae: mericarp/carpophore)
lvs opposite, often basally connate
buds with scales; w often monosymmetric
K persistent in fruit
secoiridoids
early
sympetaly
w small
embryo
short
G
inferior
lvs toothed, sec. veins palmate
dioecious, w unisexual, T small to lacking
ellagic acid
often tendrillar vines; lvs often divided and with glandular teeth
A epipetalous, ovules 2/carpel; raphides, pearl glands
berries
woody
stylodia free
hypanthium with nectary
cork origin deep-seated
endosperm lacking
resinous, lignans
/
neolignans, harman alkaloids
woody; dioecious
w small, C valvate, G unilocular
fr indehiscent
iridoids (aucubin), gutta
mostly woody; lvs if veins strong, proceed to apex of teeth
w mostly K5, persisting, mostly A∞, G mostly slightly connate
seeds often with aril; fr usu. follicles
pollen tricolpate
protandry common
w K/C/P opp A
laments rather narrow
nodes 3:3
stomata anomocytic
microsporogenesis
simultaneous
features as in
“Early Angiosperms”
borAGinAles
Boraginaceae Codonaceae Coldeniaceae Cordiaceae Ehretiaceae
(+ Lennoaceae)
Heliotropiaceae
Hydrophyllaceae Namaceae Wellstediaceae
woody
lvs stomata cyclocytic, petiole bundles annular
fr indehiscent (more or less eshy)
calcium oxalate as crystals
lvs roughly hairy; nodes unilacunar
in scorpioid
mostly 4 ovules
isokestose, higher inulins, pyrrolizidine alkaloids
A=C
polyandry
rare
AcorAles
AlismAtAles
DioscoreAles
pAnDAnAles
liliAles
ArecAles
poAles
commelinAles
ZinGiberAles
AspArAGAles
m
onocots
commeliniDs
Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae
Colchicaceae Liliaceae Petermanniaceae Smilacaceae
Bromeliaceae Eriocaulaceae Poaceae
Restionaceae
Xyridaceae
Cyperaceae Juncaceae Rapateaceae Typhaceae
(incl.
Sparganiaceae
)
Arecaceae Dasypogonaceae
Commelinaceae Haemodoraceae Hanguanaceae
Philydraceae Pontederiaceae
Cannaceae Heliconiaceae Marantaceae
Strelitziaceae
Costaceae Lowiaceae
Musaceae Zingiberaceae
Amaryllidaceae (incl. Agapanthaceae, Alliaceae) Hypoxidaceae Iridaceae
Asparagaceae (incl. Agavaceae, Hyacinthaceae, Ruscaceae) Lanariaceae Orchidaceae
Tecophilaeaceae Asphodelaceae (incl. Xanthorrhoeaceae, Hemerocallidaceae)
Cyclanthaceae Pandanaceae Triuridaceae Velloziaceae
Alismataceae Butomaceae Posidoniaceae Scheuchzeriaceae
Aponogetonaceae Hydrocharitaceae Potamogetonaceae Toeldiaceae
Araceae Juncaginaceae Ruppiaceae Zosteraceae
Acoraceae
scattered bundles in stem
no secondary thickening
mostly herbaceous
pollen monosulcate
sieve tube plastids with
protein crystals
sympodial branching
lvs parallel-veined, entire
no glandular teeth
w pentacyclic
P 3-merous, A opp. P
laments narrow
anthers broadly attached
septal nectary
single cotyledon
radicle not persistent
stem-borne roots numerous
in spadix with spathe; lvs axils with mucilaginous intravaginal squamules
ovules atropous, seeds with epidermal perisperm and copious endosperm
ethereal oils
mostly herbs and aquatics; rhizomatous
in ± scapose; G often apocarpous
placentation often laminar; endosperm helobial
often geophytes
fr capsule or berry
seed coat obliterated or with phytomelan
geophytes (bulbs, tubers, rhizomes); leaf bases often not sheathing
w T sometimes spotted, nectaries on T, anthers extrorse
seeds ∞
phytomelan lacking; fructans in stems, chelidonic acid
twining vines (often); lvs often reticulate
ovary often inferior, style short, branched
steroidal sapogenins
/
alkaloids
some woody (with terminally tufted lvs)
in
sometimes with
spathe + spadix
woody, often monopodial
lvs often palmately or pinnately pseudocompound, reduplicate-plicate
intense primary growth, large apical meristem, in often with spathe
alkaloids
herbs (mostly); lvs grassy; epidermis siliceous
mycorrhiza – (in most)
w often anemophilous, minute, chaffy, without nectaries
in thyrsus of scorpioid cymes
w monosymmetric or not, stamens few fertile
phenylphenalenones
herbs, rhizomatous, large-leafed; pseudostem common
w irregular/monosymmetric, septal nectaries
A often strongly modied/reduced, G inferior; seeds often arillate
silicic acid
UV-uorescing cell walls
(ferulic/coumaric acids)
silicic acid in leaves
cuticular waxes often in rodlets
aggregated into scallops
cerAtophyllAles
Ceratophyllaceae
aquatic; herbaceous; monoecious
lvs whorled, no pellucid dots; vessels lacking
w T0 or 9–10*, A1, G1, 1 apical ovule/carpel
pollen inaperturate, pollen tube branched, hydrophilous
woody
vessels absent
eustele
sieve tube plastids
with starch grains
lvs simple, persistent, entire
w strobilar, perfect, parts free
P parts varying, often in threes,
weakly differentiated
anthers tetrasporangiate
stamen with broad laments
pollen monosulcate
G apocarpous
(style short in most)
compitum (if present) extragynoecial
nectaries absent
siphonogamy
double fertilization
>
endosperm
embryo very small
in
cymose
fr drupe
ellagic acid
lacking
nodes trilacunar
sesquiterpenes
benzylisoquinoline alkaloids
embryo sac 8-nucleate
endosperm triploid
fr
usually
1–few-seeded
Chloranthaceae
lvs opposite, interpetiolar stipules
nodes swollen
w small T0–3, A1–5, G1, ovule 1 (apical)/carpel
sApinDAles
mostly woody (silica/silicied)
lvs often alternate, odd-pinnately compound
w often imperfect, intrastaminal disk
ethereal oils
huerteAles
Dipentodontaceae Gerrardinaceae
Petenaeaceae
Tapisciaceae
vessel elements with scalariform perforations; mucilage cells
lvs margins toothed, stipules cauline
w small, A = and opposite K, ovules 1-2/carpel
G
r
A
D
e
pArAcryphiAles
Paracryphiaceae
bruniAles
Bruniaceae Columelliaceae (incl. Desfontainia)
escAlloniAles
Escalloniaceae
mostly woody
in racemose, C free
anthers basixed, nectary disk
woody; evergreen
nodes 1:1
w polysymmetric, anthers basixed
woody
in racemose, w 4-merous, laments stout
capsule septicidal
picrAmniAles
Picramniaceae
trees, dioecious, lvs compound
extraoral nectaries, staminate w: A = and opposite C
bark bitter, anthraquinones
embryo
large
m
A
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A
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l
A
m
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A
m
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A
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A
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l
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A
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G
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Angiosperm Phylogeny Poster
Theodor C. H. Cole, Dipl. Biol.
Hartmut H. Hilger, Prof. Dr.
Dahlem Centre of Plant Sciences (DCPS)
Institute of Biology – Botany
Freie Universität Berlin
Altensteinstr. 6
D-14195 Berlin, Germany
Peter F. Stevens, Ph.D.
Missouri Botanical Garden (MoBot)
St. Louis, MO 63166-0299, USA
and
University of Missouri
–
St. Louis
Department of Biology
St. Louis, MO 63121-
4499, USA
Plant
Phylogeny
Posters
on
ResearchGate
chlorAnthAles
metteniusAles
Metteniusaceae
icAcinAles
Oncothecaceae Icacinaceae
trop. trees; few shared features; nodes usu. trilacunar
alkaloids, iridoids
Metteniusa:
lvs alternate; w bisexual, fragrant; anthers sagittate; pollen sacs moniliform, dehisce longitudinally
in axillary cyme; G(5) unilocular;
NE Andes, cloud montane forest
trop. trees; lvs exstipulate/entire; nodes often unilacunar
pedicels articulated; style short; embryo long
endosperm copious
indole and quinoline alkaloids (camptothecin)
cAryophyllAles
berberiDopsiDAles
Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae
Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae
(Chenopodiaceae) Droseraceae Nyctaginaceae Simmondsiaceae
Basellaceae Drosophyllaceae Phytolaccaceae Talinaceae
Cactaceae Frankeniaceae Plumbaginaceae
Tamaricaceae
Aextoxicaceae Berberidopsidaceae
proteAles
rAnunculAles
trochoDenDrAles
buxAles
Buxaceae (incl. Haptanthaceae)
Trochodendraceae
Nelumbonaceae
Platanaceae
Proteaceae Sabiaceae
Berberidaceae Eupteleaceae Menispermaceae Ranunculaceae
Circaeasteraceae Lardizabalaceae
Papaveraceae
lvs often divided
w parts whorled, P single or multiple whorls
G apocarpous/paracarpous, superior
berberines
mostly woody
w T often 4-merous
A epitepalous, connectives sometimes with apical appendage
woody; vessels lacking
w T 0, A∞, G>5 laterally connate with abaxial nectaries
fr aggregate of follicles
mostly woody; lvs evergreen, stomata cyclocytic
mostly monoecious, w unisexual, T ± uniform or missing
pregnane pseudoalkaloids
recepta-
cular
nectary
common
benzylisoquinolines
vAhliAles
Vahliaceae
herbaceous; lvs opposite, glandular-pubescent
w < 5 mm across, K valvate, G(3) inf, nectary
fr septicidal capsule,
K/C/A
persistent; seeds minute
Africa to India; iridoids
A
n
A
© Cole, Hilger, Stevens 2019/4 (CC-BY)
Anacardiaceae Burseraceae Kirkiaceae Nitrariaceae
Sapindaceae
Biebersteiniaceae
Meliaceae Rutaceae
Simaroubaceae
Altingiaceae Cynomoriaceae Haloragaceae
Peridiscaceae
Cercidiphyllaceae
Daphniphyllaceae
Hamamelidaceae
Paeoniaceae
Crassulaceae Grossulariaceae Iteaceae Saxifragaceae
extraoral nectaries
ethereal
oils
not in
idioblasts
w 5-merous
parts whorled
K + C free
diplostemonous
pollen tricolporate
G connate, style +
endosperm nuclear
ellagic
and
gallic acids
seed
exo-
testal
embryo
long
Ca oxalate
raphides
endosperm
nuclear
helobial
p
e
n
t
A
p
e
t
A
l
A
e
C
forming
distinct
tube
A
epipetalous
1/1/13/6/80
1/1/1-5
absent
stipules
N x
r
o
s
i
D
s
s
u
p
e
r
r
o
s
i
D
s
f
A
b
i
D
s
embryo large
endosperm scanty
COM clade
s
u
p
e
r
A
s
t
e
r
i
D
s
Flowering Plant Systematics
cycADs
GinkGo
epheDrA
W
elWitschia
Gnetum
conifers
ferns
(incl.
horsetAils
)
seeD
plAnts
l
ycophytes
AnA GrADe
AsteriDs
mAGnoliiDs
monocots
Gymnosperms
AnGiosperms
fAbiDs
mAlviDs
lAmiiDs
cAmpAnuliDs
hornworts
mosses
liverworts
rosiDs
t
rAcheophytes
COLE TCH, HILGER HH, STEVENS PF (2019) Angiosperm Phylogeny Poster (APP)
• hypothetical tree based on molecular phylogenetic (chiey plastid) data
• phylogeny, classication, and features chiey follow APweb, APG, and PPA (
Plastid Phylogenomic Angiosperm
see Li HT et al. 2019)
• 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
• branch lengths deliberate, not expressing actual time scale
• the characters listed do not necessarily apply to all members of a clade
• position of various characters on the tree uncertain
• 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
* Ceratophyllum: the alleged tepals could be bracts (issue being addressed in ongoing research and debate)
This poster is available in various languages on ResearchGate
References: Stevens PF (2022) APweb – www.mobot.org/MOBOT/research/APweb
APG IV (2016); Judd W et al. (2016); Li HT et al. (2019); Simpson M (2019); Soltis DE et al. (2014); Watson/Dallwitz (2018)
Acknowledgements to our contributors, translators, and consultants: Rubina Abid, Julien B. Bachelier, Przemysław Baranow, Zoltán Barina, Fernanda Antunes Carvalho, Christoph Dobeš†,
Johannes Enroth, Ray F. Evert, Mohamed Fennane, Vladimir N. Godin, Marc Gottschling, Zigmantas Gudžinskas, Michal Hroneš, Akitoshi Iwamoto, Chen-Kun Jiang, Anna Kagiampaki,
Kent Kainulainen, Elizabeth A. Kellogg, Sangtae Kim, Aslı Doğru-Koca, Nikos Krigas, Sanjay Kumar, Ľuboš Majeský, Pavol Márton, Diego Medan, Sergei L. Mosyakin, So Mursidawati,
Dashzeveg Nyambayar, Anastasiya V. Odintsova, Richard G. Olmstead, Batlai Oyuntsetseg, Peter H. Raven, Yasaman Salmaki, Federico Selvi, Paramjit Singh, Douglas E. Soltis, Phetlasy Souladeth,
Stoyan Stoyanov, Bach The Tran, Pramote Triboun, Zoya M. Tsymbalyuk, Orzimat T. Turginov, Magsar Urgamal, Vladimir Vladimirov, Maximilian Weigend, Michael Wink, Shahin Zarre
Actinidiaceae
Ericaceae Pentaphylacaceae
Sarraceniaceae
Balsaminaceae
Fouquieriaceae Polemoniaceae
Styracaceae
Clethraceae Lecythidaceae Primulaceae
Symplocaceae
Diapensiaceae Marcgraviaceae Roridulaceae
Theaceae
Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae
3/5/1001/4/752/10/1254/17/41707/91/28586/128/3140
1/1/2-414/166/47855/21/~1000
5/36/1345
10/67/155814/1122/36265
1/192/~260014/997/188755/68/8128/92/~2500
7/199/~45004/85/17501/2/2
1/7/1202/2/~501/11/~36015/112/2600
1/14/8502/24/345
2/94/1355
36/716/160654/762/~204007/109/~30007/33/1175
2(5)/17/~900
9/380/13000
7/12/661/4/50
9/479/6570
4/6/29
10/338/6000
18/405/~5000
2/3/4
37/749/116206/51/59022/346/12000
2/22/~200
1/11/55
2/3/181/1/8
6-11/150/3120
5/21/536
11/1743/268701/10/131
2/14/797/494/5489
1/3/38
2/46/1090 9/263/~8000
Burmanniaceae Dioscoreaceae
Nartheciaceae
Taccaceae Thismiaceae
petrosAviAles
Petrosaviaceae
stem with ring of bundles
fr follicle
East Asia
1/2/3
GentiAnAles
Apocynaceae (incl. Asclepiadaceae) Loganiaceae
Gentianaceae Gelsemiaceae
Rubiaceae
lvs opposite
interpetiolar stipules or nodal line/ridge, colleters
w corolla convolute in bud
indole alkaloids, iridoids
5/1121/20145
lAmiAles
lvs opposite; nodes 1:1
w mostly monosymmetric
A often 2(+2); gland-headed hairs with radially arranged cells
6-oxygenated avones, oligosaccharides cornoside, verbascoside (acetoside)
Acanthaceae Lamiaceae Orobanchaceae
Plantaginaceae
Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae
Byblidaceae Martyniaceae Pedaliaceae
Stilbaceae
Calceolariaceae Gesneriaceae Oleaceae Phrymaceae Verbenaceae
24/1059/~23800
woody; (semi-)parasites; mycorrhiza absent; lvs margins entire
w A epipetalous; perianth often simple, valvate, persisting
carpels/ovaries/ovules often reduced, placentation free-central; fr drupe, one-seeded
polyacetylenes, triterpene sapogenins, silicic acid
sAntAlAles
Balanophoraceae Misodendraceae Opiliaceae Schoepaceae
Loranthaceae Olacaceae Santalaceae Viscaceae
14/151/~2000
solAnAles
Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae)
Hydroleaceae Montiniaceae
Sphenocleaceae
lvs spiral, simple; nodes unilacunar
w petals plicate
K persistent in fruit
alkaloids diverse, no iridoids
5/165/~4000
oxAliDAles
Brunelliaceae Connaraceae Elaeocarpaceae
Oxalidaceae
Cephalotaceae Cunoniaceae Huaceae
lvs often compound, pulvini (sleep movement)
w A5 or multiple, branched style common
mucilage cells
oxalates
7/60/1845
ethereal oils in spherical idioblasts
(pellucid dots)
tension wood +