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Order 121. SOLANALES

Herbs (erect or climbing), shrubs, or trees, sometimes

parasitic. Indumentum of diverse types of trichomes,

often stellate or otherwise branched. Intraxylar phloem

usually present in stem and petioles. Vessels with sim-

ple perforations. Fibers mostly with simple pores.

Rays heterogeneous to homogeneous. Axial paren-

chyma apotracheal or less often scanty paratracheal.

Sieve-element plastids of S-type. Nodes unilacunar.

Leaves alternate, or rarely opposite, simple or some-

times compound, sometimes succulent, rarely scale-

like, estipulate. Stomata of diverse types. Flowers in

various types of inﬂ orescences (cymose or derived

from the cymose type) or solitary and axillary, usually

bisexual, commonly 5-merous, actinomorphic to more

or less zygomorphic. Calyx synsepalous, usually

5-lobed, persistent. Corolla sympetalous, usually

5-lobed, lobes mostly plicate in bud, contorted, imbri-

cate, or valvate. Stamens usually ﬁ ve, attached to the

corolla tube and alternate with its lobes, but in

Solanaceae-Salpiglossideae one or even three of them

transformed into staminodia; anthers usually introrse,

tetrasporangiate, opening longitudinally or by apical

pores. Tapetum secretory. Microsporogenesis simulta-

neous. Pollen grains 2-celled or sometimes 3-celled,

3-colporate, 6-polycolpate or pantoporate. Well-

developed intrastaminal nectary disc usually present.

Gynoecium usually of two, sometimes ﬁ ve, rarely

three united carpels; style usually terminal or some-

times gynobasic, with usually 2-lobed or sometimes

peltate stigma; ovary superior, (1)2(3–5)-locular, with

one or more often several or numerous ovules in each

locule or on each placenta. Ovules anatropous,

hemitropous, campylotropous, or amphitropous, uni-

tegmic, commonly tenuinucellate, usually with

endothelium. Female gametophyte of Polygonum- or

Allium-types. Endosperm cellular, sometimes nuclear,

or helobial. Fruits of diverse types, mostly baccate or

septicidal capsules. Seeds exotestal, with straight or

curved or sometimes spiral embryo and usually copi-

ous endosperm. Commonly producing various kinds

of alkaloids, ﬂ avonol and ﬂ avone glycosides, cou-

marins, acylated anthocyanins, and caffeic acid; iridoid

compound absent, n = 7–15.

Derived from the Rubiales, probably from the lower

Rubiales. The origin from the Rubiales is conﬁ rmed by

both the ﬂ ower and seed structure as well as by the

presence of intraxylary phloem.

Key to Families

1 Tall tree.

2 Secretory cavities present. Fibers with numerous

small bordered pits. Rays closely spaced, hetero-

geneous, mostly uniseriate. Axial parenchyma

occasionally diffuse, but most commonly in apo-

tracheal sheaths 1–3 cells wide or in vasicentric

sheaths 1–3 cells wide. Internal phloem present.

Leaves alternate, simple, entire. Flowers in few-

ﬂ owered terminal or subterminal cymes, bisex-

ual, actinomorphic, 5-merous except for the

gynoecium. Calyx imbricate, persistent, tubular,

with terminal teeth. Corolla funnelform, with

long tube and short, very imbricate lobes, green-

ish-white. Stamens ﬁ ve, alternating with the

corolla members; ﬁ laments attached to lower

part of corolla tube; anthers exserted, oblong,

530 Subclass VIII. LAMIIDAE

introrse, basally deeply sagittate, with a thick

connective, opening longitudinally. Pollen grains

with striate exine (Hunziker 2001). Well-

developed intrastaminal nectary disc present

around the base of ovary. Gynoecium of two car-

pels, more or less immersed in disc; style termi-

nal, elongate, with dilated, shortly 2-lobed

stigma; ovary 2-locular, but only one locule fer-

tile; ovule solitary, anatropous. Fruits large shin-

ing bright orange drupes; endocarp surrounded

by a though persistent ﬁ brous mesocarp; 2-locu-

lar, with more or less straight sterile locule

and U-shaped fertile; seed U-shaped, with curved

(U-shaped) embryo, cotyledons small; endosperm

scanty, oily. . . . . . . . . . 4. duckeodendraceae.

2 Secretory cavities absent. Stems without internal

phloem. Leaves alternate, leathery, simple, entire.

Stomata paracytic. Flowers solitary, axillary, bi-

bracteolate, 5-merous. Floral receptacle develop-

ing a short and thick gynophore. Hypogynous disc

absent. Flowers solitary, axillary, zygomorphic.

Sepals persistent, much imbricate, petals con-

torted and unilaterally induplicate, campanulate,

with short lobes. unequal but not bilabiate,

densely adpressed-ferrugineous-pubescent exter-

nally. Stamens ﬁ ve, inserted near the base of the

corolla tube; anthers long exserted, basiﬁ xed.

Gynoecium of two carpels; style one, deﬂ exed

in bud, apical; stigmas small, truncate; ovary

2-locular, stipitate. Ovules 30–50 per locule,

anatropous. Placentation axile. Fruit baccate,

crustaceous, 1–4-seeded. Endosperm copious.

. . . . . . . . . . . . . . . . . . . . . . . 9. humbertiaceae.

1 Herbs, shrubs, vines, rarely small trees.

3 Stems with internal phloem.

4 Articulated latex canals or latex cells absent.

5 Seeds with endosperm.

6 Leaves alternate or alternate to opposite.

7 Fruits mostly berries or capsules. Trees,

shrubs, perennial to annual herbs, and

lianas, prickly or not.

8 One sepal lies in the upper median

plane of the ﬂ owers. Small trees,

shrubs, perennial to annual herbs,

and lianas, often prickly, resinous, or

not resinous. Trichomes simple,

glandular, stellate, echinoid or

peltate. Cork superﬁ cial or deep-

seated, or pericycle. Plants with or

without crystal sand. Nodes with 2–3

traces. Leaves alternate or alternate

to opposite (usually alternate below,

but often becoming opposite towards

the inﬂ orescence), sometimes leath-

ery or modiﬁ ed into spines, entire,

simple to pinnately compound or

3-foliolate, rarely gland-dotted

(e.g. Anthocercis). Stomata anomo-

cytic or anisocytic. Flowers in termi-

nal or axillary inﬂ orescences, or

solitary, occasionally leaf-opposite,

small, to medium sized, usually acti-

nomorphic, mostly bisexual, rarely

unisexual (monoecious or rarely

dioecious, e.g. sometimes in Solanum

and Symonanthus). Calyx 5-lobed or

-parted, sometimes 4- or 6–7-lobed,

often enlarging in fruit. Corolla

rotated to tubular, with (4-)5(-7) pli-

cate and sometimes also contorted or

rarely merely contorted, imbricate or

valvate lobes, actinomorphic to (some

Salpiglossis) distinctly zygomorphic.

Androecium exclusively of fertile sta-

mens or rarely including staminodia

(Salpiglossideae). Stamens usually

ﬁ ve, rarely four or even (Schizanthus)

only two fertile; inserted near the base

of the corolla tube, or midway down

the corolla tube, or in the throat of the

corolla tube; anthers dorsiﬁ xed or

basiﬁ xed, extrorse (e.g. Anthocercis)

or introrse, opening longitudinally or

by apical pores or short slits. Pollen

grains 2-celled, (2)3–5(6)-colpate or

colporate, or colporoidate, or rugate,

sometimes inaperturate, rarely (spe-

cies of Salpiglossis, Bouchetia,

Nierembergia, Reyesia) in tetrads.

Gynoecium mostly of two carpels

generally oriented obliquely to the

median plane of the ﬂ ower or rarely

three carpels; style terminal, with

mostly 2-lobed stigma; ovary 2-locu-

lar [but sometimes complicated by

secondary divisions; locules second-

arily divided by ‘false septa’ (Datureae

and Nicandreae), or without ‘false

Superorder LAMIANAE 531

septa’], or 1-locular apically (as in

Capsicum). Ovules (1-) more or less

numerous on often thickened placen-

tas, anatropous or hemianatropous.

Endosperm haustoria usually present.

Fruits capsules (commonly septicidal,

or loculicidal, or valvular, or circum-

scissile – Hyoscyamus), of berries, or

drupes. Seeds with straight or curved

embryo, two semi-cylindric cotyle-

dons, and mostly oily or proteina-

ceous, rarely starchy endosperm, is

some Datura species seeds with

elaiosomes. Containing alkaloids,

arthroquinones, ursolic acid; ﬂ a-

vonols (kaempferol and quercetin)

and saponins present or absent,

n = 7–12 +.. . . . . . . . 1. solanaceae.

8 The upper median plane passes

more or less between two adaxial

sepals. Annual to perennial herbs,

or shrubs to 2 m tall (Hydrolea

spinosa). Leaves alternate, entire.

Flowers in short terminal raceme of

cymes, 5-merous. Calyx basally

connate. Anthers versatile, bas-

iﬁ xed. Nectary disc annular, ﬁ ve-

humped. Gynoecium of 2–4 carpels;

styles 3–5, glandular-pubescent;

stigma slightly tunneliform or capi-

tate; ovary 2(-3)-locular, ovules

numerous (200–300), on thickened,

axile placentae. Endosperm cellular,

with branching micropylar and cha-

lazal haustoria. Fruits globose or

ovoid, septicidal or irregularly

4-valved. Seeds longitudinally

ridged and ruminate, exotestal cells

thin-walled, endotestal cells tannin-

iferous, with a cuticle; n = (9)10

(12). . . . . . . . . .6. hydroleaceae.

7 Fruits schizocarps. Herbs or shrublets,

sometimes diffuse or prostrate, com-

monly more or less succulent. Plants

with “crystal sand”. Leaves alternate

to opposite (alternate below, those

towards the inﬂ orescence sometimes

in alternate pairs of unequal members

towards the same side of the stem),

sessile or petiolate, simple, entire, ﬂ at or

terrete, leathery or ﬂ eshy, sometimes

small and ericoid. Stomata anisocytic.

Flowers axillary, subsessile or pedicel-

late, solitary, actinomorphic or some-

what zygomorphic, 5-merous. Calyx

tubular-campanulate, lobes imbricate,

rarely valvate, persistent. Corolla acti-

nomorphic to obscurely bilabiate, pli-

cate (between the lobes). Stamens

inserted low in the corolla tube, unequal,

three longer than the other two; anthers

ovoid or oblong, introrse. Pollen grains

2-celled. Gynoecium of (3-)5 carpels;

the carpels of Alona are united to form a

conventional 5-locular ovary and style

terminal; in Nolana style gynobasic,

with peltate or capitate stigma; ovary

superior, divided into mericarps or

5-locular. Locules consist of 3–30

locelli, each locules divided horizontally

into one-ovulate locelli, or not horizon-

tally divided (Watson and Dallwitz

1992), with 1–7 ascending or appendic-

ulate, hemianatropous ovules in each,

uniseriate or in superimposed series.

Fruits separating into small 1–7-seeded

5–25 mericarps provided with germina-

tion plugs formed by the funicule, meri-

carps ﬁ ve, or 10–30 (in 1–3 series).

Seeds with curved or spiral embryo, two

cotyledons, and copious, oily endosperm.

Contain ﬂ avonols (kaempferol and quer-

cetin), n = 12.. . . . . . . 2. nolanaceae.

6 Leaves opposite, more or less entire, thick

and ﬂ eshy, usually asymmetrical. Stomata

diacytic. Plants with “crystal sand”.

Flowers solitary, sessile, in the axils.

Calyx asymmetrical, spinescent,

unequally 5-lobed, with short tube, per-

sistent. Corolla slightly zygomorphic,

conduplicate-contorted. Stamens ﬁ ve,

attached at different levels to the corolla

tube; anthers opening longitudinally.

Pollen grains 3-colporate, reticulate, colpi

with granulate membrane, tectum thick

and perforate, columellae well deﬁ ned

and some of them bifurcate. Gynoecium

of two carpels; style terminal, with simple

532 Subclass VIII. LAMIIDAE

papillate stigma; ovary 2-locular, inferior,

with the septum oblique in relation to the

median plane of the ﬂ ower. Ovules 2–3,

apotropous, pendulous from the upper

part of the locules. Fruits dry, indehiscent,

2-locular, usually 2-seeded, with thin

membranous pericarp, usually concres-

cent with the stem. Seeds with endosperm

and straight or more or less curved

embryo, n = 12. 3. sclerophylacaceae.

5 Seeds without endosperm. Shrubs or small

trees with many branches. Young parts and

ﬂ owers often densely ferrugineo-veluti-

nous. Fibers with bordered pits. Rays 1–3

cells wide, composed primarily of procum-

bent cells with upright cells only at the tips

of multiseriate rays and in uniseriate rays.

Axial parenchyma diffuse plus diffuse-in-

aggregate plus narrow bands. Leaves alter-

nate, leathery, simple, entire, coriaceous,

closely parallel-ascending, striate-veined,

midrib sometimes fusiform-thickened.

Stomata anomocytic. Flowers solitary and

extra-axillary or in few-ﬂ owered, axillary

racemes or fascicles, bisexual, actinomor-

phic or zygomorphic, 4–6-merous. Sepals

connate or rarely almost free, valvate, per-

sistent. Corolla infundibulate or campanu-

late, 4–6-lobed, lobes mostly short, equal or

unequal, valvate. Stamens 4–6, epipetalous;

ﬁ laments long-exserted, equal or unequal;

anthers exserted, versatile, sagittate, dorsi-

ﬁ xed near base. Pollen grains 3-colporate,

with thick perforate tectum and prominent

columellae, echinate. Nectary disc usually

large, ﬂ eshy, lobed. Gynoecium of 1–2 car-

pels; style terminal, long, with 2-lobed or

capitate stigma; ovary 1–2-locular, with

two ovules per locule. Fruits baccate

(drupoid?), ﬂ eshy, or chartaceous,

1–2-seeded. Seeds with plicate testa;

embryo straight, well differentiated and

large, cotyledons two or sometimes four.

The berry of Espadaea edible, with the ﬂ a-

vour of apricot. . . . . . . . . 5. goetzeaceae.

4 Articulated latex canals or latex cells usually

present, either scattered or in vertical rows, are

fairly widespread in both leaf and stem.

Autotrophic plants: herbs, shrubs, or lianas,

rarely small trees, usually with twining stems,

occasionally prostrate or creeping or erect,

sometimes succulent; some thorny xerophytes,

some with tuberous roots or stems, others rhi-

zomatous. Convolvulus, Ipomoea, Rivea twin-

ing anticlockwise. Hairs both glandular and

eglandular. Stem with intra xylary phloem.

Leaves alternate, usually petiolate, simple, cor-

date, hastate, or sagittate, entire to variously

divided, sometimes with extraﬂ oral nectaries.

Stomata usually paracytic. Flowers in terminale

cymes or solitary, bracteate, often bracteolate,

bisexual (except Hildebrandtia – plants dioe-

cious), actinomorphic, 5-merous. Hypogynous

disc present. Sepals imbricate, persistent;

Corolla valvate and plicate, or contorted and

plicate, tubular or campanulate, or urceolate,

without scale staminodia. Stamens ﬁ ve, inserted

near the base of the corolla tube, or midway

down the corolla tube. Pollen grains 2-, or

3-celled, 3-colpate to pantoporate. Intrastaminal

nectary disc usually present, annular or copular.

Gynoecium of 2–5 carpels. Sty1es one or two,

without an indusium, often ﬁ liform, simple or

forked, rarely very short or absent, stigma entire

or 2-lobed, rarely 3-lobed or stigmas 2–4, of

various shapes, globular or ellipsoid to ﬁ liform,

sometimes aplanate, rarely peltate, reniform,

conical, or funnel-shaped. Ovules anatropous,

with the micropyle directed downward and out-

ward, unitegmic, with a massive integument,

tenuninucellate or sometimes crassinu cellate.

Placentation basal. Fruits usually loculicidal

capsules, sometimes circumscissile or irregu-

larly dehiscing, rarely baccate or nutlike. Seeds

conspicuously hairy, or not; embryo large,

green, straight or curved, with folded emargin-

ate or biﬁ d cotyledons, embedded in a hard,

often cartilaginous endosperm. Often produc-

ing alkaloids of the indole and other groups,

sometimes ﬂ avonols (kaempferol and

quercetin), n = 7, 14, 15. 7. convolvulaceae.

3 Stems without internal phloem. Herbaceous root-

less (the normal root system is short-lived) para-

sites with slender, often ﬁ liform, twining, yellow,

or reddish stem, attaching on aerial parts of the

host by haustoria. Hairs when present mostly uni-

cellular or bicellular, not glandular. Intraxylary

phloem absent. Leaves alternate, reduced to

Superorder LAMIANAE 533

minute scales. Stem without intra xylary phloem.

Flowers small, 3–5-merous. Calyx imbricate.

Corolla valvate, with crenulate or ﬁ mbriate scale-

like ﬁ ve staminodia inside. Stamens ﬁ ve, inserted

in the throat of the corolla tube. Pollen grains

3-celled, 3–6-aperturate, colpate. Nectary disc

commonly present around the base of the ovary.

Gynoecium of two carpels. Stylodia two, free or

united into a simple style, stigmas globose, sub-

globose, or elongated. Ovary 2-locular; ovules

two per locule, ascending, tenuinucellate, uniteg-

mic, with a massive integument. Fruits capsules,

opening circumscissilely, irregularly, or indehis-

cent and somewhat ﬂ eshy. Embryo slender, ﬁ li-

form, green, nearly or quite acotyledonous, but

sometimes with an enlargement at one end, periph-

eral and strongly curved or spirally wound around

the starchy, oily endosperm. Present proanthocya-

nidins, ﬂ avonols (kaempferol and quercetin), n

mostly = 7, sometimes 15. . . . . .8. cuscutaceae.

1. SOLANACEAE

A.L. de Jussieu 1789 (including Atropaceae Martynov

1820, Cestraceae Schlechtendal 1833, Daturaceae

Raﬁ nesque 1828, Hyoscyamaceae Vest 1818,

Lyciaceae Raﬁ nesque 1840, Nicotianaceae Martynov

1820, Salpiglossidaceae Hutchinson 1969). 92/2300.

Subcosmopolitan, but especially diversiﬁ ed in South

America.

Classiﬁ cation based on Hunziker (2001).

1.1 BROWALLIOIDEAE (CESTROIDEAE)

Cork superﬁ cial or deep-seated; bordered pits present;

pericyclic ﬁ bers present. Corolla often zygomorphic.

Stamens inserted high in the corolla tube. Fruits usu-

ally capsular. Seeds prismatic to subglobose, with

straight or slightly curved embryo. Alkaloids mostly

pyridinic and tropane types, also steroidal, quinolinic

and phenilethylaminic types (Hunziker 2001), n =

7–14, rarely 12. – cestreae: Cestrum, Vestia, Sessea

(including Sesseopsis); metternichieae:

Metternichia; latueae: Latua; nicotianeae:

Nicotiana, Petunia (including Calibrachoa), Fabiana,

Nierembergia, Bouchetia, Leptoglossis, Hunzikeria,

Plowmania; benthamielleae: Benthamiella,

Pantacantha, Combera; francisceae: Brunfelsia;

brow allieae: Browallia, Streptosolen; schwenc-

kieae: Schwenckia, Melananthus, Protoschwenckia,

Heteranthia.

1.2 JUANULLOIDEAE

Corolla actinomorphic or zygomorphic. Stigma usu-

ally saddle-shaped. Fruits generally baccate, multi-

seeded. Seeds reniform, boomerang-shaped,

endosperm scanty, n = 12. – Juanulloa, Dyssochroma,

Ec tozoma, Hawkesiophyton, Markea, Merinthopodium,

Rahowardiana, Schultesianthus, Trianaea.

1.3 SOLANOIDEAE

Corolla actinomorphic, aestivation varied: imbricate,

vavate, contorted. Stamens ﬁ ve, inserted low in the

corolla tube. Stigma small, unmodiﬁ ed. Fruits usually

baccate. Seeds ﬂ attened, discoid, with coiled embryo,

endosperm usually abundant, n = 10, 12, 14, 17. – nica-

ndreae: Nicandra; mandragoreae: Mandragora;

datureae

: Datura, Brugmansia; lycieae: Lyceum,

Phrodus, Grabowskia; solaneae: Witheringia,

Brachistus, Cuatresia, Deprea, Discopodium, Exode-

conus, Jaltomata, Nothocestrum, Physalis (including

Margaranthus), Tzeltalia, Quincula, Leucophisalis

(Physaliastrum), Chamaesaracha, Oryctes, Tubo-

capsicum, Capsicum, Aureliana, Athenaea, Darcya-

nthus, Larnax, Withania (including Mellisia,

Archiphysalis), Solanum, Cyphomandra, Lycopersicon,

Lycianthes, Triguera, Normania; iochrominae:

Iochroma, Saracha, Acnistus, Eriolarynx, Vassobia,

Dunalia; atropeae: Atropa (including Pauia);

jaboroseae: Jaborosa, Salpichroa, Nectouxia; solan-

dreae: Solandra; Hyoscyameae: Hyoscyamus,

Anisodus, Atropanthe, Physochlaina, Przewalskia,

Scopolia.

1.4 SALPIGLOSSOIDEAE

Corolla actinomorphic or zygomorphic, imbricate.

Pollen grains in monads or in tetrads. Capsules

septicide-loculicidal, with two or four valves, many

seeded (up to 300 in Salpiglossis, or 2–25 in Reyesia).

Endosperm cellular. Present pyridinic alkaloids,

n = 11. – Salpiglossis, Reyesia.

1.5 SCHIZANTHOIDEAE

Cork pericycle; pericycle ﬁ bers absent. Flowers zygo-

morphic, stalk curved and the two really upper petals

form the lower lip which is 3–4-lobed, and the lowest

petal forms the simple or slightly 2-lobed upper lip.

Stamens four: two fertile and two staminodial;