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544 Subclass VIII. LAMIIDAE

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Superorder LAMIANAE 545

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Order 123. OLEALES

Trees or shrubs, sometimes woody climbers or scram-

blers; Jasminum twining anticlockwise. The indumen-

tum characteristically consists of scale-like, peltate

trichomes, sometimes glandular or appearing as trans-

parent or sunken dots in the leaf; groups of secretory

type of trichomes frequently form extraﬂ oral nectaries

that are common in some of the genera. Small acicular

or prismatic crystals of calcium oxalate occur in both

stem and leaf. Vessels with scalariform or scalariform

and simple perforations (in Forsythia infrequently

occur scalariform perforations, and vestigial scalari-

form plates present in Syringa and Fontanesia); pits

vestured, lateral pitting alternate, often more or less

distinctly vestured (Baas et al. 1988). Fibers with bor-

dered or simple pits, sometimes septate. Rays heteroge-

neous to homogeneous. Axial parenchyma apotracheally

diffuse or more often paratracheal. Sieve-element plas-

tids of S-type. Nodes unilacunar. Leaves often decidu-

ous, or evergreen, opposite or rarely (some species of

Jasminum) alternate, petiolate, simple and often entire

or lobed to pinnately compound or trifoliolate or unifo-

liolate, conduplicate (Chionanthus), estipulate. Stomata

usually anomocytic. Domatia occurring in six genera

and numerous species and manifested as pits, or pock-

ets, or hair tufts. The nuclei of mesophyll parenchyma

cells contain characteristic crystalline inclusions.

Flowers mostly rather small, basically in dichasial

cymes but usually modiﬁ ed so as to give the appear-

ance of racemes, panicles or fascicles, sometimes soli-

tary, bisexual or rarely polygamous or dioecious,

actinomorphic, usually 4-merous. Calyx mostly small,

4(-15)-lobed, valvate, rarely absent (some spp. of

Fraxinus). Corolla sympetalous, 4(-15)-lobed, but

sometimes, as in spp. of Fraxinus very deeply lobed or

divided and seemingly choripetalous, lobes imbricate,

valvate (or induplicate-valvate), or contorted, some-

times (as in Forestiera and some spp. of Fraxinus)

obsolete, rarely corolla absent. Stamens usually two,

rarely four (Hesperelaea, Tessarandra), very rarely

three or ﬁ ve, attached to the corolla tube (when the tube

is expressed); ﬁ lantherous, or with sessile anthers;

546 Subclass VIII. LAMIIDAE

anthers dorsiﬁ xed or basiﬁ xed, introrse, tetrasporan-

giate, with locules usually back to back, often apicu-

late, opening longitudinally; endothecium developing

ﬁ brous thickenings. Tapetum secretory. Microsporo-

genesis simultaneous. Pollen grains 2-celled or very

rarely 3-celled, tricolpate, or tricolporate, occasionally

rupate, with reticulate ornamentation. A nectary

intrastaminal disc sometimes present around the base

of the ovary. Gynoecium of two carpels, mostly median;

style terminal, with entire, 2-lobed or biﬁ d stigma or

sometimes stigma sessile; ovary superior, 2-locular,

with two, one or four to many ovules per locule,

attached to the apex, side, or base of the partition.

Ovules pendulous or ascending, with dorsal raphe,

usually anatropous, unitegmic, with usually massive

integument, tenuinucellate, with persistent endothelium

around the lower part of female gametophyte. Female

gametophyte monosporic, usually of Polygonum-type,

rarely (Olea) bisporic of Allium-type. Endosperm

cellular or sometimes (Jasminum) nuclear. Fruits of

various types: usually 1-seeded berries, drupes, some-

times samaras (Fraxinus), woody loculicidal or

circumscissile (Menodora) capsule. Seeds small, exot-

estal (exotesta palisade or not, endotesta ﬁ brous),

embryo straight, cotyledons ﬂ at, endosperm copious,

oily, thick-walled, or cotyledons storage and endosperm

almost absent. Usually producing mannitol, ﬂ avones

and ﬂ avonols, carboxycyclic iridoid compounds,

myricetin, and the phenolic glycosides orobanchin and

syringin as well as verbascosides (Jensen et al. 2002),

ursolic acid. Triterpenes and saponins are also present.

n = 11, 13–14, 18, 20, 22–24.

Some authors, including Dahlgren, consider Oleaceae

a separate order Oleales related to the Rubiales

(Gentianales), especially to the Loganiaceae, and others

(Cronquist 1981, 1988) placed them in the

Scrophulariales, and Thorne (1992a, b, 2000, 2006) – in

the Lamiales. Wood anatomy supports the treatment of

Oleaceae in or near the Rubiales, closest to the

Loganiaceae s. 1. (Baas et al. 1988: 168). According to

Hegnauer (1969), chemical evidence also supports

afﬁ nities of the Oleaceae with Rubiales. However, recent

molecular data (rbcL sequences, Olmstead et al. [1992])

and restriction site mapping of the chloroplast DNA

(Downie and Palmer 1992) seem to agree with the posi-

tion of the Oleaceae near the Scrophulariaceae (Jensen

1992). “It may have been the presence of seco-iridoids,

which never occur in Lamianae, that was decisive for

Dahlgren’s removal of Oleaceae from the superorder.

However, the type of seco-iridoids in Oleaceae is unique

when compared to those found in other seco-iridoid-

containing taxa.… Thus, the presence of seco-iridoids

alone may not be justiﬁ cation for keeping Oleaceae in

Gentianales” (Jensen 1992: 293). Serological studies

support afﬁ nities with both Rubiales and Scrophulariales

(Piechura and Fairbrothers 1979). On the other hand,

the chloroplast DNA data suggest that the Oleaceae are

more closely related to the Scrophulariales than to the

Rubiales (Downie and Palmer 1992). In some respects

the Oleales occupy an intermediate position between

Gentianales and Lamiales assemblage.

1. OLEACEAE

Hoffmannsegg et Link 1809 (including Bolivariaceae

Grisebach 1838, Forestieraceae Endlicher 1841,

Fraxinaceae Vest 1818, Jasminaceae A.L. de Jussieu

1789, Ligustraceae G. Meyer 1836, Nyctanthaceae J.

Agardh 1858, Schreberaceae Schnitzlein 1857–1870,

Syringaceae Horaninow 1847). 25/615–900. Nearly

cosmopolitan, but best developed in Southeast Asia

and Australia; the largest genus Jasminum (450) is

conﬁ ned to the Old World, mainly to tropical and sub-

tropical regions.

jasmineae: Menodora, Jasminum; forsythieae:

Abeliophyllum, Forsythia; fontanesieae: Fontanesia;

myxopyreae: Myxopyrum, Nyctanthes, Dimetra;

fraxineae: Fraxinus; schrebereae: Comoranthus,

Schrebera; oleeae: Syringa (including Ligustrina),

Ligustrum, Olea, Chionanthus, Forestiera, Haenian-

thus, Tessarandra, Noronhia, Notelaea, Nestegis,

Osmanthus, Phyllyrea, Picconia, Hesperelaea.

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Order 124. LAMIALES

Herbs and subshrubs, less often shrubs and trees,

sometimes lianous. Young twigs often quadrangular.

Stems sometimes with anomalous structure. Vessels

with simple or rarely scalariform perforations; lateral

pitting alternate. Fibers usually with simple pits, often

septate. Rays mostly heterogeneous of various types

to homogeneous. Axial parenchyma paratracheal, very

scanty or absent. Intraxylary phloem rare. Sieve-

element plastids usually of S-type, but in Acanthaceae

also of Pcs- and Pc-types. Nodes unilacunar or rarely

trilacunar or multilacunar. Leaves alternate or oppo-

site, sometimes verticillate, simple or sometimes com-

pound, almost always estipulate; leaf mesophyll cells

sometimes contain oil bodies. Stomata of various

types. Flowers in cymose or racemose inﬂ orescences

or solitary, bisexual or rarely unisexual, usually zygo-

morphic. Perianth 5-merous or less often 4-merous.

Sepals more or less connate, lobed or toothed, some-

times bilabiate, sometimes persistent. Corolla sympet-

alous, often bilabiate, sometimes more or less

actinomorphic, lobes imbricate. Stamens attached to

the corolla tube and alternate with the corolla lobes, in

ﬂ owers with 4-merous perianth as many as the lobes or

only two, in ﬂ owers with 5-merous perianth usually

only four fertile stamens and the upper (adaxial) sterile

or wanting or staminodial, or only two fertile stamens

and the others are wanting or staminodial. Anthers tet-

rasporangiate or less often disporangiate, opening lon-

gitudinally. Tapetum secretory or rarely (Bignoniaceae)

amoeboid. Microsporogenesis simultaneous. Pollen

grains 2-celled or 3-celled, 3-aperturate or of various

548 Subclass VIII. LAMIIDAE

derived types. Intrastaminal nectary disc usually pres-

ent. Gynoecium mostly of two united median carpels,

sometimes pseudomonomerous, rarely of three or four

carpels; style terminal or gynobasic, with 2-lobed

stigma (style sometimes very short and the stigma is

essentially sessile); ovary superior or less often semi-

inferior or inferior, with (1)2 or mostly many ovules

per locule. Ovules anatropous, hemitropous, amphitro-

pous, or less often orthotropous, unitegmic, tenuinu-

cellate, usually with endothelium. Female gameto-

phyte usually of Polygonum-type. Endosperm cellular.

Endosperm haustoria developed. Fruits capsules or

capsulelike, or sometimes indehiscent and then drupa-

ceous or baccate, sometimes separating into drupelike

mericarps. Seeds exotestal, with copious endosperm or

endosperm scanty or wanting. Usually producing vari-

ous iridoid compounds (but not secoiridoids) and the

phenolic glycoside orobanchin, ﬂ avones, but only

rarely alkaloids.

Lamiales are related to the Oleales. In some respects

the Buddlejaceae are a connecting link between these

orders.

Key to Families

1 Flowers essentially actinomorphic.

2 Perianth usually 4-merous.

3 Trees or shrubs, rarely lianas or suffrutescent

herbs, often with lepidote or stellate or branch-

ing trichomes and/or glandular hairs. Vessels

with simple perforation, without vestured pits.

Fibers with simple or obscurely bordered pits,

septate (in some Buddleja, Peltanthera,

Nuxia). Rays heterogeneous. Axial paren-

chyma sparse, paratracheal, or absent. Leaves

opposite or less often verticillate, rarely (a few

spp. of Buddleja) alternate, entire or more

often dentate or lobed, with interpetiolar

stipules usually represented by a line, rarely

well developed. Stomata paracytic, anomo-

cytic, or anisocytic, and these types often

mixed on one plant (Oxelman, Kornhall and

Norman 2004). Flowers in various types of

inﬂ orescences, bisexual or often functionally

dioecious, bracteate and bracteolate, 4-mer-

ous, rarely (Peltanthera) 5-merous. Calyx

lobed or very deeply cleft, persistent, imbri-

cate. Corolla with imbricate or valvate lobes,

tubular, funnelform. Stamens four, inserted

midway down the corolla tube, or in the throat

of the corolla tube; anthers bilocular, or 4-loc-

ular (Androya), dorsiﬁ xed or peltate, introrse.

Pollen grains 2-celled, 3(4)-colporate. Nectary

disc absent, except for Gomphostigma.

Gynoecium of two carpels, or of four carpels

(in some Buddleja species); style with a capi-

tate or lobed stigma; ovary 2-, or 4-locular,

superior, locules secondarily divided by ‘false

septa’, or without ‘false septa’; ovules numer-

ous, hemitropous, pendulous or ascending, on

thickened, axile placentas. Fruits septicidal

capsules, rarely barriers (Nicodemia). Seeds

often winged, sometimes with pockets between

the exotesta and the endothelium (Norman

2000); embryo straight, small to large,

endosperm oily or starchy, copious to scanty.

Produce iridoids aucubin and catalpol, and

verbascosides (Jensen 2000). n = 7, 14, 15,

19. . . . . . . . . . . . . . . . . . . . 1. buddlejaceae.

3 Small herbs, terrestrial, or subaquatic to

aquatic.

4 Fruits capsular, with numerous seeds. Small

per ennial, rarely annual herbs arising from

a central taproot with numerous sympodial

branches; branches procumbent to ascend-

ing, ﬂ uted, often with four sharp ﬂ anges

and 2- or 4-rounded ridges. Leaves oppo-

site, sessile, connate, simple, entire, linear,

slightly scabrid along the margins and stem

ridges, stipulate or estipulate, stipules, if

present, interpetiolar. Young stems tetrago-

nal. Flowers axillary, solitary, bracteolate

(the bracteoles sepal-like), small, sessile.

Calyx divided almost to the base, persistent.

Corolla campanulate, imbricate, with a ring

of hairs in the throat. Stamens usually four,

included, ﬁ laments about as long as the

anthers, alternating with the corolla mem-

bers; anthers 2-locular, introrse. Pollen grains

3-colporate. Gynoe cium of two carpels; style

short, persistent with a capitate stigma; ovary

2-locular, partly inferior. Ovules 20–50 per

locule, anatropous, on peltate placentae that

are attached to the base of the septum.

Endosperm cellular. Endosperm haustoria

chalazal or micropylar. Fruits loculicidal and

valvular capsules, laterally compressed, obo-

void to ovoid. Seeds small, wingless, cuboi-

dal, with a smooth surface, translucent;

Superorder LAMIANAE 549

embryo straight; endosperm copious.

Detected verbascosides and cornoside,

n = 11. . . . . . . . . . . . . . 2. polypremaceae.

4 Fruits separate into four basally attached,

1-seeded, setulose, dry mericarps. Mostly

subaquatic to aquatic, succulent, small

creeping herbs with prostrate rooting stems

emitting short, leafy branches, bearing

essential oils. Leaves opposite, leathery,

with ﬂ at connate petioles, ciliate toward

base, minutely denticulate distally, more or

less ﬂ eshy, entire, simple, obscurely gland-

dotted, aromatic. Flowers solitary, terminal

or axillary, minute. Sepals connate, imbri-

cate, persistent. Petals connate, imbricate,

corolla subrotate, without a ring of hairs in

the throat. Stamens four, inserted to the

corolla tube, alternating with the corolla

members; anthers 2-locular, dorsiﬁ xed,

introrse. Pollen grains 3- colporate, rhom-

boidal, exine regularly dimorphic, the polar

regions and borders of colpi psilate and

tectate- perforate, the mesocolpal regions

intectate and baculate. Gynoecium of two

carpels, with long or short, slender gynoba-

sic style and inconspicuous stigma; ovary

superior, 4-locular (ostensibly), or 2-locular

with four locelae via false septa, with one

erect anatropous ovule per locule; placenta-

tion basal. Mericarps comprising nutlets.

Seeds with copious endosperm. Contain

verbascoside. . . . 3. tetrachondraceae.

2 Flowers apetalous.

5 Gynoecium of two united carpels; style

gynobasic, deeply cleft or often divided to the

base. Slender perennial or annual herbs; some

of them entirely submerged aquatics with

underwater pollination, others are amphibi-

ous, and few are terrestrial in wet habitats with

aerial pollination. Glandular, fan-like scale

hairs in the leaf axils, in terrestrial and amphib-

ious species additionally peltate, disc-like

hairs. Vascular system of the stem very

reduced, xylem with or without vessels.

Leaves small, opposite, often in rosettes at

branch tips, usually linear and biﬁ d when sub-

merged, linear to spatulate when ﬂ oating and

aerial, epulvinate. Stomata present, sporadi-

cally on submerged leaves, mainly conﬁ ned to

one surface (adaxial, when ﬂ oating), or on

both surfaces. Flowers without perianth, min-

ute, unisexual, monoecious or dioecious, usu-

ally solitary in axils or rarely one ﬂ ower of

each sex in the same axil, usually with two,

minute, hornlike bracteoles. Male ﬂ ower con-

sists of one stamen, rarely of 2–3 stamens;

anthers reniform. Pollen grains small, spheroi-

dal, intectate, with three, not well-deﬁ ned

apertures or inaperturate, 3-coplate. Ovary

superior, somewhat compressed, laterally

4-lobed, apically somewhat indented; each

carpel longitudinally divided by an intrusive

partition from the carpellary midrib into two

locelli each with one pendulous, anatropous

ovule with ventral raphe. Endosperm with ter-

minal, chalazal and micropylar (the latter the

more aggressive) haustoria. Fruits dry, 4-lobed,

with each lobe winged or keeled, splitting into

four one-seeded mericarps. Seeds minute,

with membranous testa, straight or slightly

curved embryo and ﬂ eshy, oily endosperm;

cotyledons short. Contains carbocyclic iridoid

aucubin (Wieffering 1966), verbascosides, ﬂ a-

vones and caffeic acid, n = 3–6, rarely 10, 14,

19–20. . . . . . . . . . . . . . 13. callitrichaceae

5 Gynoecium monocarpellate (derived from a

pseudomonomerous one), with long subulate

stylodium stigmatic throughout. Perennial

water herbs with creeping, sympodial rhizome

and erect stems, whose upper parts usually

project above the water. Vascular system of

the axial organs very reduced. Sieve-element

plastids of S-type. Leaves verticillate, entire,

linear, estipulate, provided with minute, decid-

uous, peltate, glandular trichomes; submersed

leaves long, pale green and ﬂ accid, the emer-

gent leaves obovate to linear, dark green and

rigid. Flowers small and inconspicuous, soli-

tary and sessile in the axils of upper leaves,

bisexual or sometimes some or all of them

unisexual, actinomorphic, apetalous, anemo-

philous. Calyx reduced to 2–4-lobed or suben-

tire rim around the top of the ovary. Stamens

solitary, abaxial, with slender ﬁ lament and

large, red, tetrasporangiate, introrse anther

opening longitudinally. Tapetum secretory.

Micro sporogenesis simultaneous. Pollen

grains 3-celled, 4–6-colporate. Ovary inferior,

550 Subclass VIII. LAMIIDAE

1-locular, with one apical and pendulous, ana-

tropous, unitegmic to ategmic, tenuinucellate

ovule with an endothelium. Endosperm cellu-

lar, with inconspicuous terminal haustoria.

Female gametophyte of Polygonum-type.

Fruits drupaceous, with very thin, ﬂ eshy exo-

carp and hard endocarp. Seeds with large,

elongate, straight embryo and very thin, pro-

teinaceous endosperm. Producing the carbo-

cyclic iridoid aucubin and catalpol, n = 8, 15,

16. . . . . . . . . . . . . . . . . . .14. hippuridaceae.

1 Flowers mostly more or less zygomorphic. Perianth

mostly 5-lobed.

6 Plants with ericoid habit and densely crowded,

verticillate leaves and glandular trichomes. Trees,

small shrubs or shrublets; shrubs single-stemmed,

or multistemmed, young branches often angular

and puberulous. Vessels simple, rarely scalari-

form. Axial parenchyma vasicentric, scarce or

lacking. Leaves opposite or rarely alternate,

entire, narrowly lanceolate to linear, with recurved

margins, usually estipulate. Inﬂ orescences termi-

nal or rarely on axillary branches (Retzia). Flowers

bisexual, each ﬂ ower subtended by leafy bract

and two bracteoles. Calyx of 4–5 sepals, usually

tubular basally, with 4–5 free, imbricate to valvate

lobes. Corolla actinomorphic or zygomorphic,

with narrowly funnel-shaped tube and 5- or less

often 4-lobed, imbricate. Stamens usually four or

ﬁ ve, equal; ﬁ laments free, short to long, slender

or stout, inserted in the throat of the corolla tube,

or between the lobes; anthers 2-locular, dorsi-

ﬁ xed. Pollen grains 3-colporate, with punctate

tectum and granular membrane of colpi. Small,

inconspicuous nectary disc usually present.

Gynoecium of two carpels; ovary superior, 2-loc-

ular, often becoming 1-locular by abortation or

loss of septum. Ovules 1–4(-6) in each locule;

ovules anatropous, ascending. Style terminal, ﬁ li-

form, erect, stigma capitate or minutely bilobed.

Fruits loculicidal and valvular capsules. Seeds

reticulate, with straight embryo and oily or starchy

endosperm. Present C-8–C-11 iridioid glucosides

(Frederiksen et al. 1999), and verbascosides;

n = 12 (Retzia), 19 (Nuxia). . . . . .5. stilbaceae.

6 Plants not ericoid. Vessels with simple perforations

(except for a few members of Bignoniaceae).

7 Plants not insectivorous. Placentation not free-

central.

8 Seeds mostly with copious endosperm

(except some Gesneriaceae).

9 Corolla not scarious. Flowers mostly

entomophilous or ornithophilous.

10 Leaves mostly alternate.

11 Ovules (2-) more or less numer-

ous on each placenta. Herbs, sub-

shrubs, or shrubs, sometimes

climbers, rarely trees (Paulownia,

Wightia); plants sometimes

succulent, rarely hydrophytic

(Limnophila), sometimes totally

parasitic, or partially parasitic, or

autotrophic. Often provided with

hairs that commonly have a basal

cystolith, often also with other

types of trichomes including glan-

dular hairs. Vessels with simple

perforation, without vestured pits;

wood partially storied or not; axial

parenchyma apotracheal or para-

tracheal, usually very sparse, or

absent. Nodes unilacunar. Leaves

alternate or less often opposite,

rarely verticillate, entire or some-

times dissected, often reduced to

scales, or absent (e.g. Lathraea).

Stomata usually anomocytic or

anysocytic. Flowers in various

types of cymose or racemose inﬂ o-

rescences or sometimes solitary,

bisexual, usually zygomorphic.

Calyx deeply (1–2)4–5-lobed,

persistent, lobes imbricate or val-

vate, rarely open. Corolla (4)5(-8)-

lobed, imbricate or valvate, often

bilabiate, sometimes spurred, per-

sistent or deciduous, rarely (spp.

of Besseya) wanting. Stamens usu-

ally four (with adaxial one stamin-

odial or staminodium absent) or

sometimes ﬁ ve (as in Verbascum)

or only 2(3) (with the abaxial pair

reduced or wanting); ﬁ laments

sometimes appendiculate; anthers

bilocular, or rarely (Selagineae)

unilocular, tetrasporangiate or

rarely disporangiate, opening by

longitudinally. Pollen grains

Superorder LAMIANAE 551

2-celled, (2)3(-6)-colporate or

panto-colpate, rarely inaperturate,

mostly more or less reticulate.

A unilateral or annular intrastami-

nal nectary disc usually present.

Gynoecium of two or rarely three

carpels; style with 2(-4)-lobed to

capitate or crateriform stigma;

ovary 2(3)-locular, rarely

(Orobanchoideae) 1-locular, with

more or less numerous (rarely sol-

itary), pendulous to ascending,

anatropous or hemitropous, sel-

dom amphitropous or campylotro-

pous ovules in each locule. Fruits

usually capsules, septicidal or less

often loculicidal, or occasionally

poricidal, or circumscissile, or

splitting irregularly, rarely bac-

cate, very rarely schizocarps.

Seeds small to minute, exotestal,

mostly angular, winged or wing-

less, without amyloid; embryo

straight or slightly curved, in para-

sitic taxa usually undifferentiated;

endosperm oily, very rarely

(Monttea) lacking. Contain iridoid

glycosides (especially aucubin and

catapol), pyrrolizidine alkaloids,

ﬂ avonoids and ﬂ avones, saponins.

In Digitalis containing glycosides

(digitalin and lanoxin) and lacking

iridoids (Fischer 2004), n = 6–9,

11, 13, 15–17, 20, 21, 23–28.

. . . . . . . . .6. scrophulariaceae.

11 Ovules usually solitary in locule.

Small shrubs, subshrubs, or rarely

perennial herbs. Imperforate trac-

heary elements are all tracheids.

Axial parenchyma scanty, vasi-

centric, less commonly diffuse.

Leaves alternate, simple, entire,

coriaceous, estipulate. Flowers in

heads (Globularia) or spikes

(Poskea) with or without involu-

cre of bracts, bisexual, zygomor-

phic. Calyx persistent, tubular,

5-lobed, actinomorphic or rarely

slightly bilabiate. Corolla 4–5-

lobed, bilabiate (the upper lip of

two petals more or less shorter than

the 3-petaled lower one), lobes

imbricate. Stamens four, inserted

at the top of the corolla tube;

anthers reniform, at ﬁ rst 2-locular,

at length the loculi conﬂ uent and

opening by a single slit. Pollen

grains 3(4)-colporate, tectate-per-

forate (Globularia) or semitectate

(Poskea), with granular to spinulose

colpus membranes. Intrastaminal

nectary disc annular, semicircular,

or glandlike or absent. Gynoecium

pseudomonomerous, with a slen-

der style and capitate or shortly

2-lobed stigma; ovary superior,

1-locular, with anatropous, pendu-

lous ovule. Fruits nutlike, envel-

oped by the persistent calyx. Seeds

with straight long embryo sur-

rounded by scanty endosperm.

Present iridoids: globularin, catal-

pol, globularicosin, globularidin,

asperulin, and lytanthosalin

(Hegnauer 1989b), n = 8, 16.

. . . . . . . . . 11. globulariaceae.

10 Leaves mostly opposite.

12 Ovary bilocular.

13 Ovary superior to semi-inferior.

Annual or perennial herbs or

shrubs (Stemotria), glandular

pubescent to densely villose,

rarely glabrous. Stems erect,

prostrate or ascending. Leaves

cauline or in a basal rosette,

sessile to petiolate, linear lan-

ceolate to ovoid-orbicular,

acuminate, entire or pinnatiﬁ d

to pinnatisect. Inﬂ orescence

thyrsic, with accessory ﬂ owers,

paired, each pair composed of

a terminal ﬂ ower and the ﬂ ower

from its reduced prophyll.

Flowers distinctly zygomor-

phic, 4-merous, bright yellow;

sepals valvate, free to the base,

persistent. Corolla with whorl

tube, usually with shoe-like

552 Subclass VIII. LAMIIDAE

invagination of lower lit, with

trichome nectaries. Stamens

two or three (Stemotria trian-

dra), ﬁ laments short, inserted

in a distinct, dark purple ring at

the base of the corolla tube;

anthers 2-locular. Gynoecium

of two carpels; style terminal,

with inconspicuous or slightly

capitate stigma. Capsule septi-

cidal and loculicidal. Seeds

numerous, testa with anticlinal

walls sinuous (straight);

endosperm present, n = (8)9.

. . . . . . .4. calceolariaceae.

13 Ovary inferior. Perennial herbs

or subshrubs. Leaves opposite,

sometimes oblique, membra-

nous, serrate or irregularly

dentate, estipulate, petioles

connected by a raised line.

Stomata anomocytic. Flowers

in dense terminal or axillary

cymes, bisexual, slightly zygo-

morphic or nearly actinomor-

phic. Calyx adnate to the ovary,

with four or ﬁ ve more or less

unequal lobes, open, persistent.

Corolla campanulate or funnel-

shaped, lobes imbricate or ind-

uplicate-valvate. Stamens two,

epipetalous; ﬁ laments short;

anthers linear- oblong, some-

times latrorse, cohering round

style. Pollen grains 3-colpate

(Silvianthus) or 5–6-colpate

(Carlemannia) intectate,

broadly reticulate. Nectary disc

conspicuous, shortly cylindri-

cal or conical. Gynoecium of

two carpels; style elongate, with

clavate or fusiform, biﬁ d

stigma; ovary 2-locular, with

many (30–100) ovules on axile

or subbasal placenta. Fruits

loculicidal capsules

(Carlemannia) or fruits open

by ﬁ ve irregular, longitudinal

valves (Silvianthus), calyx per-

sistent. Seeds many, exotesta

cells narrow, radial walls

thickened, endothelium persis-

tent (Silvianthus) or polygo-

nal, all walls thickened

(Carlemannia); endosperm

ruminate (Silvianthus), more or

less ﬂ eshy, copious; embryo

small. 10. carlemanniaceae.

12 Ovary 1-locular, or with two pari-

etal placentas. Herbs or sub-

shrubs, rarely shrubs or small

trees, sometimes lianas or epi-

phytes. Usually provided with

uniseriate hairs or stalked glands

or both. Secretory cavities pres-

ent or absent, when present – with

oil, or with resin. Fibers with sim-

ple pits and septate. Axial paren-

chyma paratracheal, or very

sparse, or absent. Nodes unilacu-

nar, trilacunar, or pentalacunar.

Leaves mostly opposite, rarely

verticillate or alternate, entire or

dentate, rarely pinnatiﬁ d, com-

monly estipulate. Stomata usually

anisocytic (often very large).

Flowers in various types of

cymose or seldom racemose

inﬂ orescences or solitary in the

leaf axils, sometimes epiphyllous,

bisexual or rarely monoecious,

mostly zygomorphic. Sepals ﬁ ve,

free or usually connate into a

lobed tube, valvate or rarely

imbricate. Corolla 5-lobed, usu-

ally bilabiate and often spurred,

sometimes nearly or quite actino-

morphic, lobes imbricate, adaxial

(posterior) ones usually internal.

Stamens commonly four, or two,

with the anthers connivent in

pairs or all together (posterior

reduced or absent), less often a

single pair of stamens developed,

rarely (Ramonda, some Sinningia

spp.) all ﬁ ve stamens developed;

often 1–3 staminodia present in

place of missing stamens, anthers

Superorder LAMIANAE 553

basiﬁ xed or rarely dorsiﬁ xed,

opening by longitudinal slits or

by apical (rarely basal) poles.

Pollen grains (2)3(4)-colporate.

Intrastaminal nectary disc usually

present, usually annular or cupu-

lar, sometimes unilateral or of

discrete glands. Gynoecium of

two carpels, with slender style

and usually 2-lobed stigma; ovary

superior or more or less inferior,

1-locular, or with two parietal

placentas that are usually more or

less intruded and bifurcate, some-

times more or less joined and

dividing the ovary into two or

even four chambers. Ovules

numerous, anatropous or very

rarely orthotropous. Fruits usu-

ally loculicidal or rarely septi-

cidal, or rarely valvular capsules,

rarely berries. Seeds small, with

straight embryo and oily

endosperm (Gesnerideae) or no

endosperm (Cyrtandroideae).

Iridoids absent. Contain ﬂ avonoids,

anthocyanins (pelargonidin, malvi-

din, cyanidin, gesnerin, delphini-

din, and etc.), cornosids and some

genera also verbascosides and

orobanchine, n = (4)8–18(64); n =

30 + in Coronanthereae (Smith

1996).. . . . . . . . .8. gesneriaceae.

9 Corolla scarious. Perennial or annual

herbs, rarely small shrubs or subshrubs,

terrestrial to aquatic, sometimes succu-

lent. Trichomes non-glandular or glandu-

lar with a head of one or a few cells.

Fibers very short, with simple or bor-

dered pits. Nodes unilacunar, trilacunar,

or multilacunar. Leaves usually all basal

and alternate, rarely cauline and alternate

or opposite, mostly sheathing at the base,

with more or less curvi-palmate or nearly

parallel venation, or leaves much reduced.

Stomata mostly diacytic, sometimes ano-

mocytic. Flowers small, mostly in brac-

teate spikes or heads, or solitary, without

bracteoles, usually bisexual, but monoe-

cious in Littorella and gynomonoecious

in Bougueria, actinomorphic, anemo-

philous or partly entomophilous, some-

times cleistogamous, 4-merous or rarely

3-mcrous. Calyx membranous, lobed or

cleft, sometimes abaxial segments more

or less connate, imbricate. Corolla

4-lobed or 4-dentate, rarely 3-lobed

(Bougueria), lobes imbricate. Stamens

usually four or rarely 1–2 (Bougueria);

ﬁ laments usually very long; anthers long-

exerted, dorsiﬁ xed, versatile. Pollen

grains 2–3-celled, 4–14-porate.

Gynoecium of two carpels, with slender

style and usually 2-lobed stigma; ovary

superior, 2-locular, with 5–50, usually

anatropous to hemitropous ovules in each

locule, or (Bougueria and Littorella)

1-locular with usually one basal ovule.

Fruits membranous circumscissile cap-

sule (Plantago) or nutlike in persistent

calyx. Seeds with straight or (Bougueria)

curved embryo in copious translucent

endosperm. Present aucubin and related

iridoid glucosides, phenylethanoid gly-

cosides (Ronsted et al. 2003), alkaloids,

terpenes and saponins, n = 4–6, 12.

. . . . . . . . . . . . . . 12. plantaginaceae.

8 Seeds mostly with scanty endosperm

(except some Acanthaceae-

Nelsonioideae and Nesogenaceae) or

endospermless.

14 Seeds ﬂ at, often alate with mem-

branous or corky wing. Trees,

shrubs, or often woody lianas,

rarely herbaceous vines (Tourretia)

or erect perennial herbs (Argylia,

Incarvillea, Niedzwedzkia); the

twiners twining anticlockwise

(Tecoma). Vessels usually with

simple perforations, but in some

genera some of them with reticulate

or scalariform perforation plates

that have numerous bars. Fibers

with simple or bordered pits. Rays

homogeneous to sometimes hetero-

geneous. Axial parenchyma mostly

paratracheal. Nodes unilacunar,

with three to several traces. Leaves