Takhtajan Armen. Flowering Plants

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Superorder ASTERANAE 503

with long, narrow lobes. Anthers dorsiﬁ xed, mostly

calcarate and caudate. Pollen grains globose, mostly

spiny, usually excavate, sometimes honeycombed.

Style arms usually long, acute, with single stigmatic

area on inner surface; style hairs usually evently dis-

tributed along the style-branches and on the shaft

below. Cypselas with twin hairs. Pappus usually pres-

ent, sometimes heteromorphic. Plants mostly unarmed.

Laticifers often present; n = (7-)9–10(-13). – gymnar-

rheneae: Gym narrhena; moquinieae: Moquinia,

Pseudostifftia; vernonieae: Vernonia, Piptocarpha,

Stokesia, Lychno phora, Elephantopus, etc.; liabeae:

Munnozia, Chrysactinium, Liabum, Oligactis, etc.;

cichorieae (lactuceae): Scolymus, Cichorium,

Tolpis, Amoseris, Andryala, Hieracium, Catananche,

Krigia, Malaco thrix, Stephanomeria, Hyoseris,

Hypochaeris, Leonto don, Picris, Urospermum,

Hedypnois, Rhagadiolus, Scorzonera, Epilasia,

Tragopogon, Koelpinia, Dubyaea, Soroseris,

Prenanthes, Lactuca, Stepto rhamphus, Scariola,

Cephalorrhynchus, Mycelis, Cicerbita, Lapsana,

Crepis, Ixeris, Youngia, Taraxa cum, Chondrilla,

Heteracia, Launaea, Reichardia, Sonchus, etc.; gun-

delieae: Gundelia; arctotideae-arctotidinae:

Arctotis, Berkheya, etc.; arctotideae-gorterii-

nae: Gorteria, Gaza nia, Cullumia, Hirpicium, etc.;

arctotideae-eremothamninae: Eremothamnus,

Hoplophyllum.

5.5 ASTEROIDEAE

Perennial to annual herbs or shrubs, less often trees,

sometimes scandent, epiphytic or aquatic, sometimes

succulent. Leaves alternate or opposite, not spiny.

Florets one to many, (3–4-)5(-6)-merous; marginal

ﬂ owers often differentiated, radiate and 3-dentate or

outer ﬂ owers female, with ﬁ liform corolla, central

ﬂ owers actinomorphic and usually shallowly lobed;

anthers basiﬁ xed, mostly ecalcarate and often ecau-

date, usually slender. Pollen grains mostly spiny and

caveate, never honeycombed. Styles generally pilose

mainly at the tips of the branches, mostly with stig-

matic areas in two separate lines. Cypselas with twin

hairs. Pappus present or absent, usually of bristles or

scales or coroniform, sometimes auriculiform or of

awns, sometimes heteromorphic. Laticifers usually

absent; n = (4-)9–10(-19). – corymbieae: Corymbium;

senecioneae: Ligularia, Farfugium, Doronicum,

Tussilago, Petasites, Adenostyles, Dendrocacalia,

Paragynoxys, Tetradymia, Robinsonia, Brachyglottis,

Werneria, Gynoxys, Cineraria, Gynura, Crasso-

cephalum, Senecio, Emilia, Othonna, Lopholaena,

Kleinia, Euryops, etc.; calenduleae: Dimorphotheca,

Osteospermum, Oligocarpus, Tripteris, Calendula,

Gibbaria, Chrysanthemoides, Garuleum, etc.;

gnaphalieae: Phagnalon, Athrixia, Filago, Micropus,

Evax, Bombycilaena, Cymbolaena, Lucilia, Metalasia,

Relhania, Loricaria, Cassinia, Helichrysum, Iﬂ oga,

Lasiopogon, Gnaphalium, Antennaria, Leontopodium,

Angianthus, etc.; astereae: Grangea, Chrysopsis,

Engleria, Grindelia, Gutierrezia, Solidago, Haplopap-

pus, Chrysothamnus, Pteronia, Dichrocephala, Bellis,

Myriactis, Calotis, Callistephus, Aster, Erigeron,

Lachnophyllum, Psychrogeton, Melanodendron, Commi-

dendrum, Diplostephium, Olearia, Felicia, Micro-

glossa, Conyza, Nolletia, Chrysocoma, Baccharis,

etc.; anthemideae: Santolina, Anthemis, Achillea,

Chamaemelum, Matricaria, Chamomilla, Cladanthus,

Anacyclus, Chrysanthemum, Tanacetum, Microcephala,

Tridactylina, Cancrinia, Lepidolopha, Leucanthemum,

Cotula, Soliva, Artemisia, Lasiospermum, Erioceph-

alus, Ursinia etc.; inuleae: Inula, Pulicaria, Blumea,

Jasonia, Pegolettia, Nauplius, Telekia, Buphthalmum,

Carpesium, Amblyocarpum, Pluchea, Laggera,

Nicolasia, Pterocaulon, Epaltes, Blumeopsis, Ade-

lostigma, Karelinia, Streptoglossa, Stenachaenium,

Cylindrocline, etc.;

heliantheae: Blepharispermum,

Arnica, Bahia, Flaveria, Hymenopapus, Gaillardia,

Helenium, Hymenoxys, Dyssodia, Tagetes, Pectis,

Porophyllum, Bidens, Chrysanthellum, Coreopsis,

Cosmos, Dahlia, Fitchia, Echinacea, Rudbeckia,

Zinnia, Verbesina, Wedelia, Helianthus, Simsia,

Galinsoga, Tridax, Melampodium, Sigesbeckia,

Silphium, Ambrosia, Iva, Parthenium, Xanthium,

Adenostemma, Eupatorium, Trichogonia, Campulo-

clinium, Ayapana, Critonia, Koanophyllon, Chromo-

laena, Bartlettina, Neomirandea, Symphy opappus,

Ageratum, Stevia, Brickellia, Helogyne, Liatris,

Fleischmannia, Micania, Ageratina, etc.

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A. Takhtajan, Flowering Plants, 511

Trees, shrubs, subshrubs, and herbs. Vessels mostly

with simple perforations. Sieve-element plastids of

S-type. Nodes unilacunar, trilacunar, or multilacunar.

Leaves alternate or more often opposite, sometimes

verticillate, estipulate or with stipules. Flowers nearly

always sympetalous. Tapetum usually secretory.

Microsporogenesis simultaneous. Gynoecium mostly

of two united carpels; ovary superior to inferior. Ovules

unitegmic and usually tenuinucellate. Endosperm

nuclear or cellular, often with haustoria. Fruits of vari-

ous types. Seeds with or without endosperm.

The subclass Lamiidae is most probably derived

from the most primitive Cornanae – Desfontainiales.

According to Olmstead et al. (1992: 259; 1993), this

lineages is one of the most strongly supported clades

in the higher dicots.

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Soltis DE, PS Soltis, PK Endress, and MW Chase. 2006.

Phylogeny and evolution of Angiosperms. Sinauer,

Sanderland, MA.

Superorder LAMIANAE

Order 120. RUBIALES (GENTIANALES)

Trees, shrubs, subshrubs, and perennial or annual herbs.

Intraxylary (internal) phloem present in all investigated

members; interxylary (included) phloem of the foram-

inate type present in Antoniaceae, Strychnos and some

species of Logania. Vessels with simple perforations or

rarely scalariform perforations; lateral pitting mostly

alternate. Fibers with bordered or more often simple

pits, sometimes septate. Vestured pits present or rarely

(Gelsemium and Logania) absent. Rays heterogeneous

to homogeneous or wanting. Axial parenchyma of

diverse types or absent. Sieve-element plastids of S-type

or Ss-type. Nodes trilacunar, unilacunar, or multi-

lacunar. Leaves opposite or sometimes verticillate,

rarely alternate, simple and usually entire, estipulate or

less often with interpetiolar stipules; leaf mesophyll

cells sometimes contain oil bodies. Colleters often

present. Stomata mostly paracytic or anomocytic.

Flowers in cymose or less often racemose inﬂ ores-

cences, sometimes solitary, bisexual or rarely unisexual,

mostly actinomorphic or nearly actinomorphic, with

double perianth. Corolla sympetalous, lobes usually

contorted, less often imbricate or valvate. Stamens

usually as many as and alternate with the corolla lobes,

in a single whorl, or very rarely (Dialypetalanthus)

stamens 16–26, in two whorls, usually attached to the

corolla tube; anthers tetrasporangiate, opening longitu-

dinally or very rarely (as in Exacum) by apical pores.

Tapetum secretory. Microsporogenesis simultaneous or

(some Apocynaceae) successive. Pollen grains 2-celled

or 3-celled, mostly 3-colporate. Intrastaminal nectary

disc present or absent. Gynoecium of two or less often

3–5 (up to eight) united carpels; stylodia more or less

united into a terminal style; ovary superior to inferior,

mostly 1-locular, with (1-) many ovules in each locule

or on each placenta. Ovules anatropous to hemitropous,

unitegmic, tenuinucellate, sometimes with endothe-

lium. Female gametophyte usually of Polygonum-type.

Endosperm nuclear or rarely (Mitrasacme, Voyriella)

cellular (Johri et al. 1992), very rarely (in some

Apocynaceae) with chalazal haustorium. Fruits capsules

or of diverse other types. Seeds exotestal, with more or

less straight embryo and copious endosperm, rarely

endosperm nearly wanting. Usually producing iridoid

Subclass VIII. LAMIIDAE

512 Subclass VIII. LAMIIDAE

compounds and often also alkaloids, but without car-

diotonic glycosides.

The basal group within the Lamiidae, probably

directly connected with the Hydrangeales and have a

common origin from the saxifragalean stock.

Key to Families

1 Plant without a latex-system, and without cardio-

tonic glycosides.

2 Intraxylary (internal) phloem present.

3 Stigma twice dichotomously branched.

Shrubs, subshrubs, or lianas with simple hairs

or almost glabrous. Vessels mostly with sim-

ple perforations, except for a few scalariform

perforations, plates with few bars close to the

pith in Mostuea; intervascular pits small, ves-

tured (Mostuea) or not vestured (Gelsemium).

Fibers with small bordered pits, nonseptate

(Gelsemium) or some septate ones present

(Mostuea). Rays heterogeneous (Mostuea) or

homogeneous (Gelsemium). Axial paren-

chyma scanty vasicentric (Gelsemium) or

absent (Mostuea). Secondary thickening

anomalous from a single cambial ring. Leaves

opposite or whorled, entire or obscurely sinu-

ate-dentate; stipulate (stipeles interpetiolar to

intrapetiolar, sometimes reduced to a stipular

sheath joining the opposite leaf bases) or

estipulate. Colleters present in the axils of the

leaves, bracts, and sepals, or only in the axils

of the leaves. Flowers in axillary or terminal

cymes or sometimes solitary, bracteate, nearly

always heterostylous, actinomorphic or nearly

so, bisexual, with variously shaped bracts.

Perianth and androecium 5-merous, occasion-

ally (Mostuea) 4-merous. Sepals free or con-

nate up to the middle, equal or unequal,

imbricate, persistent. Corolla funnel-shaped,

tube much longer than the lobes, lobes imbri-

cate, equal or subequal. Stamens four, rarely

ﬁ ve, inserted near the base of the corolla tube,

or midway down the corolla tube, equal or

unequal; ﬁ lament very short; anthers dorsi-

ﬁ xed, extrorse (Gelsemium) or latrorse. Pollen

grains 3-colporate, with long, unbranched col-

umellae and granulate colpus membrane

(Gelsemium) or ﬁ nely suprastriate, with long

and branched columellae and more or less

smooth colpus membrane. Gynoecium of two

carpels; styles 1–2, free to partially joined,

stigmas two, or four (each bifurcated). Ovary

superior, 2-locular, glabrous or hairy, with two

(Mostuea) or 2–8 (Gelsemium) anatropous

ovules in each locule. Fruits septicidal

(Gelsemium) or loculicidal (Mostuea) cap-

sules. Seeds rather large, ﬂ attened, winged

(Gelsemium) or wingless, with rather thin

testa, small, straight or curved embryo and

copious, starchy (Gelsemium), or bony

(Mostuea) endosperm. Producing seco-

iridoids, C-17 indole alkaloids, quercetin,

kaempferol; n = 4, 8 (Gelsemium) or 10

(Mostuea).. . . . . . . . . . . . . 1. gelsemiaceae.

3 Stigma not branched.

4 Mostly trees and shrubs, sometimes woody

lianas or herbs.

5 Fruits capsules.

6 Seeds free (not embedded in the pulp).

7 Plants producing iridoids and small

amount of alkaloids. Shrubs, sub-

shrubs, or herbs, Logania tortuosa is

switch-plant, glabrous or with simple

hairs. Vessels usually with simple, or

rarely scalariform perforations, with-

out vestured pits. Fibers with simple

pits. Nodes unilacunar (with one to

several traces), trilacunar or multila-

cunar. Secondary thickening develop-

ing from a conventional cambial ring,

or anomalous. Leaves opposite, entire,

simple, sometimes reduced to scales,

connected at the base by more or less

distinct stipular line or a short ochrea.

Flowers in terminal or axillary cymose

inﬂ orescences or solitary, bisexual or

unisexual (dioecious), actinomorphic.

Perianth and androecium 5-merous,

rarely (only Logania micrntha)

4-merous. Sepals subequal, basally

connate, imbricate, with colleters

inside or not. Corolla rotated or cam-

panulate, lobed, lobes imbricate.

Stamens attached to the corolla tube

or corolla throat, in female ﬂ owers

smaller or absent; anthers dorsiﬁ xed,

introrse. Pollen grains 3-colporate.

Gynoecium of two carpels; style

thick, with large, subglobose, or

Superorder LAMIANAE 513

oblong stigma; ovary superior, 2-loc-

ular, with several (3–50) ovules per

locule; ovules anatropous or hemi-

anatropous. Fruits septicidal or loculi-

cidal (dehiscing only in the upper

half) capsules, or schizocarp (Watson

and Dallwitz 1992). Seeds more or

less ellipsoid, wingless, with a

honeycomblike reticulation; embryo

straight, cylindrical, endosperm

starchy, n = 16. . . . 2. loganiaceae.

7 Plants not producing either iridoids

or alkaloids. Erect or climbing shrubs

or small to large trees, glabrous or

with simple hairs. Wood diffuse

porous, not storied, secondary thick-

ening anomalous present. Vessels

with simple perforations, with ves-

tured pits; xylem with tracheids

(abundant in Usteria) or without tra-

cheids. Fibers with small bordered

pits. Leaves opposite, simple, entire

or dentate, connected at the base by

an interpetiolar line, short sheath, or

ciliolate rim; colleters often present

in the axils of leaves and bracts and

at the base of sepals. Stomata ano-

mocytic, paracytic, or encyclocytic.

Flowers in usually terminal cymes

or in panicles, actinomorphic or

(Usteria) zygomorphic, 5-merous or

(Usteria) 4-merous. Calyx some-

times (Antonia) surrounded by

numerous imbricate, scalelike bracts,

5–4-lobed or partite, lobes or seg-

ments equal or (Usteria) outer lobe

enlarged and petaloid. Corolla tubu-

lar, tube often slender; lobes ﬁ ve or

four, valvate. Hypogynous disc small

or absent. Stamens ﬁ ve, four or

(Usteria) one, inserted in the throat

of the corolla tube;. anthers dorsi-

ﬁ xed, latrorse or introrse. Pollen

grains 3-colporate. Gynoecium of

two carpels; style deciduous or per-

sistent, with capitate or shortly

2-lobed stigma; ovary superior or

rarely inferior, 2-locular, with numer-

ous (15–50), anatropous or amphi-

tropous ovules. Fruits septicidal,

2-valved capsules. Seeds winged, the

wing diaphanous, reticulate-veined;

embryo relatively large, straight, or

incurved in ﬂ eshy or starchy

endosperm. Iridoids not detected,

n = 11 (Usteria). . 4. antoniaceae.

6 Seeds embedded in pulp. Small, gla-

brous shrubs to small trees, rarely

scrambling to semiclimbing. Branches

terete, quadrangular, or 4-winged.

Wood diffuse porous; not storied.

Vessels mostly with simple perfora-

tions, rarely (a few spp. Labordia),

with scalariform perforations, with

vestured pits. Leaves opposite, petio-

late to sessile, simple, entire, stipulate

(stipules interpetiolar or intrapetiolar)

or estipulate. Colleters present or

absent. Flowers in terminal (Labordia)

or axillary, ramiﬂ orous or cauliﬂ orous

(Geniostoma) inﬂ orescences or occa-

sionally solitary, unisexual or gynodi-

oecious, actinomorphic, 5-merous.

Sepals nearly free or connate, imbri-

cate, inside mostly with colleters at the

base, margin mostly ﬁ mbriate. Corolla

campanulate to rotate, lobed, lobes

imbricate or contorted. Stamens ﬁ ve,

inserted at the corolla mouth; ﬁ laments

mostly short, not appendiculate; anthers

dorsiﬁ xed, introrse; often with a dis-

tinct apical appendage. Pollen grains

3-pororate or with more than three

pores in zonoporate or pantoporate

position, psilate or coarsely reticulate.

Gynoecium of two or three (Labordia)

carpels. Ovary superior, 2- or rarely

3-locular, with many anatropous or

amphitropous ovules in each locule;

style mostly short; stigma about as large

as the ovary, clavate, ellipsoid or sub-

globose, sometimes more or less

oblong. Capsules with thick and woody

exocarp, septicidal to subseptifragal,

with two (or three) valves breaking

away from and exposing the persistent

and pulpy placentas. Seeds wingless,

ellipsoid, ovoid, or subglobose, intruded