See

discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/299597890

Angiosperm Phylogeny Poster - Flowering

Plant Systematics, APG IV (2016) - Outdated!

Data · April 2016

DOI: 10.13140/RG.2.1.5169.5768

CITATIONS READS

0 3,710

2 authors:

Theodor C. H. Cole Hartmut H. Hilger

Freie Universität Berlin Freie Universität Berlin
174 PUBLICATIONS 31 CITATIONS 281 PUBLICATIONS 1,576 CITATIONS

SEE PROFILE SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Angiosperm Phylogeny Poster View project

Bryophyte Phylogeny Poster View project

All content following this page was uploaded by Theodor C. H. Cole on 04 April 2016.

The user has requested enhancement of the downloaded file.

 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
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

d
Austrobaileyaceae Schisandraceae (incl. Illiciaceae) Trimeniaceae
A e
tiglic acid, aromatic terpenoids

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 (incl. 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

i “Early Angiosperms”

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
p flw G apocarpous; placentation often laminal; endosperm helobial; embryo large/green Aponogetonaceae Butomaceae Posidoniaceae Scheuchzeriaceae
Araceae Hydrocharitaceae Potamogetonaceae Zosteraceae
e
Petrosaviales
stem with ring of bundles
r Monocots Ca oxalate
raphides
fr a follicle; East Asia Petrosaviaceae
m
s
atactostele (scattered bundles)
no secondary thickening
mostly herbaceous
endosperm
nuclear
helobial
often twining vines; lvs often reticulate
ovary often inferior, style short, branched; steroid sapogenins/alkaloids Dioscoreales Burmanniaceae Dioscoreaceae Nartheciaceae Taccaceae
nectaries
some woody (with terminally tufted leaves)
Pandanales
pollen monosulcate when present
woody; vessels absent
sieve tube plastids with
septal
infl spadix Cyclanthaceae Pandanaceae Velloziaceae
eustele; nodes
sieve tube plastids protein crystals
often geophytes (bulbs, tubers, rhizomes)
sympodial branching Alstroemeriaceae Corsiaceae Melanthiaceae Philesiaceae
Liliales
with starch grains flw tepals sometimes spotted, nectaries at tepals; many seeds, seeds coat (testa) cellular
lvs simple, persistent, entire vascular bundles in stem scattered
flw parts free, strobilar, perfect lvs parallel-veined, entire phytomelan lacking; fructans in stems, chelidonic acid, steroid saponins Colchicaceae Liliaceae Petermanniaceae Smilacaceae
P parts varying, often in threes, no glandular teeth
weakly differentiated flw pentacyclic, P 3-merous, A opp. P often geophytes; anomalous sec. growth Amaryllidaceae ( incl. Agapanthaceae , Alliaceae) Hypoxidaceae Iridaceae
Asparagales
stamen with broad filaments filaments narrow capsule or berry
pollen monosulcate anthers broadly attached, seed coat obliterated or with phytomelan Asparagaceae ( incl. Agavaceae, Hyacinthaceae, Ruscaceae) Lanariaceae Orchidaceae
septal nectary; single cotyledon
G apocarpous (style short in most)
nectaries absent adventitious roots Tecophilaeaceae Asphodelaceae (incl. Xanthorrhoeaceae, Hemerocallidaceae)
monopodial, woody; lvs pinnately pseudocompound, reduplicate-plicate

Arecales
embryo very small intense primary growth, large apical meristem
aromatic terpenoids infl 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 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
Cannaceae Heliconiaceae Marantaceae Strelitziaceae
Zingiberales
silicic acid in leaves G inferior, A often strongly modified/reduced
cuticular waxes often in rodlets
aggregated into scallops
seeds arillate; silicic acid Costaceae Lowiaceae Musaceae 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

Proteales
mostly woody; flw tepals often 4-merous
A epitepalous, connectives sometimes with apical appendage Nelumbonaceae Platanaceae Proteaceae Sabiaceae
woody; vessels lacking; idioblasts in cortex; flw tepals missing, A∞
G>5 laterally connate with abaxial nectaries; fr aggregate of follicles Trochodendrales Trochodendraceae
E axial

u
nectary
common
mostly woody; unisexual, dioecious; lvs evergreen, stomata cyclocytic
flw tepals ± uniform or missing; pregnane pseudoalkaloids Buxales Buxaceae (incl. Haptanthaceae)
d unisexual, dioecious; lvs toothed, sec. veins palmate
i
c
flw tepals small to lacking
ellagic acid Gunnerales Gunneraceae Myrothamnaceae
mostly woody; lvs veins proceed to apex & teeth; flw K5–∞, persisting
o
t
mostly A∞, G mostly slightly connate
seeds often with aril; fr often both ventrally/dorsally dehiscent Dilleniales Dilleniaceae
benzylisoquinolines absent

s Cynomoriaceae
lvs with glandular teeth; often hypanthium, apically unfused carpels, stigma decurrent Altingiaceae Daphniphyllaceae Hamamelidaceae
pollen tricolpate
flw K/C/P opp A
fr mostly dry, dehiscent
myricetin, flavonols Saxifragales Cercidiphyllaceae Grossulariaceae Paeoniaceae
filaments rather narrow often tendrillar vines; lvs often divided and with glandular teeth Crassulaceae Haloragaceae Saxifragaceae

Vitales
anthers basifixed A epipetalous, 2 ovules per carpel; raphides, pearl glands
nodes trilacunar
stomata anomocytic
berries Vitaceae
ethereal oils absent cork origin deep-seated

stipules
endosperm lacking
resinous, lignans/neolignans, harman alkaloids Zygophyllales Krameriaceae Zygophyllaceae
nodes
3:3 flw small, G often 3-merous, nectary: intrastaminal disk
seeds often with aril (red-orange)
infl cymose Celastrales Celastraceae (incl. Hippocrateaceae, Brexiaceae, Parnassiaceae) Lepidobotryaceae
S
u lvs often compound, pulvini (sleep movement)
Brunelliaceae Connaraceae Elaeocarpaceae
p
flw A5 or multiple, branched style common
mucilage cells; oxalates Oxalidales Cephalotaceae Cunoniaceae Huaceae
Oxalidaceae
e
r F Achariaceae Euphorbiaceae Rafflesiaceae Ochnaceae Podostemaceae

c E r a lvs margins toothed

flw G often tricarpellate Malpighiales Chrysobalanaceae
Clusiaceae
Hypericaceae
Linaceae
Passifloraceae
Phyllanthaceae


Rhizophoraceae
Salicaceae
u o b
o
d s R i flw often “papilionaceous”: wing, standard, keel, mostly G1 Erythroxylaceae Malpighiaceae Picrodendraceae Violaceae
r o
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)

o s i lvs mostly simple with stipules Barbeyaceae Elaeagnaceae Rosaceae

t d
s
N fix
flw K valvate (and hypanthium) persisting
carpels with 1 ovule, stigma dry; dihydroflavonols Rosales Cannabaceae Moraceae Ulmaceae
s embryo large Dirachmaceae Rhamnaceae Urticaceae (incl. Cecropiaceae)
endosperm scanty lvs mostly alternate
flw pentacyclic
parts alternating
K + C (free)
flw often unisexual, G mostly inferior
parietal placentation; cucurbitacins Cucurbitales Apodanthaceae Begoniaceae
Anisophyllaceae Coriariaceae
Corynocarpaceae
Cucurbitaceae
Datiscaceae
Tetramelaceae
G connate lvs mostly undivided; flw small, unisexual
(A polyandrous) anemophilous, thus T reduced or lacking, G mostly inferior
Betulaceae Fagaceae Myricaceae Rhoipteleaceae
pollen tricolporate
ellagic and gallic acids
infl spikes or catkins; fr 1-seeded, mostly nuts
ectomycorrhiza; tannins, dihydroflavonols Fagales Casuarinaceae Juglandaceae Nothofagaceae Ticodendraceae
lvs with glandular teeth
flw A obdiplostemonous, nectary on filament
stems jointed at nodes; ethereal oils, ellagic acid Geraniales Geraniaceae Francoaceae (incl. Ledocarpaceae, Melianthaceae, Vivianiaceae)
lvs opposite, colleters (glandular hair on adaxial surface of petiole base) Combretaceae Myrtaceae Penaeaceae (incl. Oliniaceae)
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 Stachyuraceae
Geissolomataceae 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 embryo
lvs margins toothed, stipules cauline
flw small, A = and opposite K, ovules 1-2/carpel Huerteales Dipentodontaceae Gerrardinaceae Petenaeaceae Tapisciaceae
s large
bark fibrous; hairs often stellate Bixaceae Malvaceae (incl. Bombacaceae, Sterculiaceae, Tiliaceae)
endo-

Malvales
flw K often valvate
sperm
scanty petals contorted, A often ∞; mucilage
Cistaceae Cytinaceae Muntingiaceae Sarcolaenaceae
cyclopropenoid fatty acids, flavones Dipterocarpaceae Neuradaceae Sphaerosepalaceae Thymelaeaceae
lvs alternate; flw often 4-merous
often clawed petals, A often many, 2×K Bataceae Caricaceae Limnanthaceae Salvadoraceae
hypogynous (often gynophore); infl racemose
myrosin cells, glucosinolates Brassicales Brassicaceae Cleomaceae Moringaceae Tovariaceae
Liverworts Capparaceae Koeberliniaceae Resedaceae Tropaeolaceae
woody; lvs stomata cyclocytic
Mosses
petiole bundles annular; fr fleshy
calcium oxalate as crystals Berberidopsidales Aextoxicaceae Berberidopsidaceae
Hornworts woody; (semi-)parasites; without mycorrhiza; lvs margins entire
flw A epipetalous; perianth often simple, valvate, persisting Balanophoraceae Misodendraceae Opiliaceae Schoepfiaceae
Lycophytes free-central pendulous placentation; fr one-seeded
polyacetylenes, triterpene sapogenins, silicic acid Santalales Loranthaceae Olacaceae Santalaceae Viscaceae
Ferns mostly herbaceous; without mycorrhiza
(incl. horsetails) Aizoaceae Caryophyllaceae Molluginaceae Polygonaceae
G often unilocular with central placentation, Amaranthaceae Didiereaceae Nepenthaceae Portulacaceae
Palmferns
Ginkgo S pollen colpate, surface spiny
betalains or anthocyanins (latter, e.g., in Caryophyllaceae) Caryophyllales [incl. Chenopodiaceae]
Basellaceae
Droseraceae
Drosophyllaceae
Nyctaginaceae
Phytolaccaceae


Simmondsiaceae
Talinaceae
Ephedra u mostly woody; lvs mostly undivided, hydathode teeth Cactaceae Frankeniaceae Plumbaginaceae Tamaricaceae
Seed Gymnosperms
Welwitschia
Gnetum p flw often 4-merous, K much smaller than C, persisting
Plants Conifers

ANA grade
e
r
intrastaminal disk, G inferior; fr drupaceous
diverse iridoids Cornales Cornaceae
Curtisiaceae
Grubbiaceae
Hydrangeaceae

Hydrostachyaceae
Loasaceae
Nyssaceae
Magnoliids a Actinidiaceae Ericaceae Polemoniaceae Sarraceniaceae
s lvs teeth, theoid; nodes unilacunar
Angiosperms Monocots
t
flw 5-merous, pentacyclic
nonhydrolyzable tannins, ellagic acid, hydroquinones Ericales Balsaminaceae
Clethraceae
Fouquieriaceae
Lecythidaceae


Primulaceae
Roridulaceae
Styracaceae
Theaceae
Fabids e
r
tropical trees; lvs exstipulate/entire; nodes often unilacunar Ebenaceae Myrsinaceae Sapotaceae Theophrastaceae
pedicels articulated; style short; embryo long
Malvids
Rosids

Lamiids
i
d
fr 1-seeded (drupe); endosperm copious
indole and quinoline alkaloids (camptothecin) Icacinales Oncothecaceae Icacinaceae
s A tropical trees; few common features; nodes usu. trilacunar
Asterids Campanulids
s 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
t woody; dioecious
e
r L
flw small, C valvate, G unilocular; fruit indehiscent
iridoids (aucubin), gutta Garryales Eucommiaceae Garryaceae (incl. Aucubaceae)
i a lvs spiral, simple; nodes unilacunar
m Convolvulaceae (incl. Cuscutaceae) Solanaceae (incl. Nolanaceae)
Theodor C. H. Cole, Dipl. Biol.
d
s ii
flw petals plicate; K persisting
diverse alkaloids, no iridoids Solanales Hydroleaceae Montiniaceae Sphenocleaceae
Institute of Pharmacy and Molecular Biotechnology (IPMB) d herbaceous; Africa to India; lvs opposite, glandular-pubescent
Heidelberg University s
Vahliales
mostly sympetalous stipules lacking; flw < 5 mm across, K valvate, G(3) inf, nectary
Im Neuenheimer Feld 364,D-69120 Heidelberg, Germany nectary gynoecial fr septicidal capsule, K/C/A persistent; seeds minute; iridoids Vahliaceae
ovules unitegmic
endosperm cellular
A=C, lvs opposite, colleters Apocynaceae (incl. Asclepiadaceae) Loganiaceae
Gentianales
iridoids common
epipetalous flw corolla convolute in bud
polyandry
rare
indole alkaloids; iridoids Gentianaceae Gelsemiaceae Rubiaceae
G(2) lvs opposite; nodes 1:1; flw mostly monosym.
A often 2(+2); ethereal oils in gland-headed hairs
Acanthaceae Lamiaceae Orobanchaceae Plantaginaceae
Lamiales
infl
cymose 6-oxygenated flavones, rosmarinic acid, Bignoniaceae Lentibulariaceae Paulowniaceae Scrophulariaceae
ellagic acid
lacking
oligosaccharides: cornoside, verbascoside (acetoside) Byblidaceae Martyniaceae Pedaliaceae Stilbaceae
lvs roughly hairy; nodes unilacunar Gesneriaceae Oleaceae Phrymaceae Verbenaceae
infl scorpioid; mostly 4 ovules

Boraginales Boraginaceae Codonaceae Coldeniaceae Cordiaceae Ehretiaceae (+ Lennoaceae)

isokestose, higher inulins,
Prof. Dr. Hartmut H. Hilger pyrrolizidine alkaloids
Dahlem Centre of Plant Sciences (DCPS)
Heliotropiaceae Hydrophyllaceae Namaceae Wellstediaceae
woody; lvs serrate
Institute of Biology – Plant Morphology and Systematics
Freie Universität Berlin early
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 sympetaly flw often monosymmetric Asteraceae Goodeniaceae Pentaphragmataceae
flw small

Asterales
________________________________________________________________________________________ plunger pollination (long style)
embryo nodes trilacunar Calyceraceae Menyanthaceae Rousseaceae
- hypothetical tree based on molecular phylogenetic data (April 2016) short
- phylogeny, classification, and features chiefly follow APweb and APG
inulin, sesquiterpenes, secoiridoids Campanulaceae (incl. Lobeliaceae) Stylidiaceae
- 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 C mostly woody; infl racemose, C free
anthers basifixed, nectary disc Escalloniales Escalloniaceae
- branch lengths deliberate, not expressing actual time scale a
m Bruniales
- the characters listed do not necessarily apply to all members of the according clade woody, evergreen; nodes 1:1
- position of various characters on the tree uncertain flw polysymmetric, anthers basifixed Bruniaceae Columelliaceae (incl. Desfontainia)
- References: Stevens PF (2016) 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 (2015) delta-intkey.com/ a woody (except Apiaceae)
Apiaceae Griseliniaceae Pennantiaceae
A piales
lvs often divided; nodes usu. multilacunar
- for family characteristics see: FGVP, Kubitzki K, ed. (1990 ff)
n G inferior infl mostly umbel; drupe or schizocarp
Special thanks for ongoing consultation since inception of this poster to Peter F. Stevens (Missouri Botanical Garden)
u (Apiaceae: mericarp/carpophore) Araliaceae Myodocarpaceae Pittosporaceae
Acknowledgements to Julien B. Bachelier, Fernanda Antunes Carvalho, Christoph Dobeš, Ray F. Evert, Mohamed Fennane, Marc Gottschling, Chen-Kun Jiang, Anna Kagiampaki, Diego Medan, fr usually
l 1–few-seeded

Paracryphiales
Sergei Mosyakin, Sofi Mursidawati, Richard G. Olmstead, Peter H. Raven, Federico Selvi, Douglas E. Soltis, Zoya M. Tsymbalyuk, Maximilian Weigend, Michael Wink, Shahin Zarre woody; infl racemose, flw 4-merous,
i filaments stout, capsule septicidal Paracryphiaceae
Angiosperm Tracheophyte Bryophyte 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

View publication stats