Takhtajan Armen. Flowering Plants

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554 Subclass VIII. LAMIIDAE

opposite or sometimes verticillate,

rarely alternate, simple or more

often pinnately compound or trifo-

liolate or palmately compound.

Stomata anomocytic, paracytic, or

diacytic (Kigelia). Inﬂ orescences

terminal to axillary, ramiﬂ orous to

cauliﬂ orous cymes, or reduced to a

single ﬂ ower. Flowers mostly

large, bracteate and bracteolate,

5-merous. Calyx campanulate,

lobed or toothed, sometimes bila-

biate (spathelike in Spathodea) or

truncate, rarely calyptrate (Lundia

spp.). Corolla campanulate or fun-

nelform, usually more or less

strongly zygomorphic, sometimes

bilabiate, lobes imbricate or rarely

valvate. Stamens inserted near the

base of the corolla tube, or mid-

way down the corolla tube, typi-

cally four in two pairs, the ﬁ fth

(adaxial) staminodial or absent,

rarely all ﬁ ve fertile (Oroxylum) or

two fertile and three staminodial

(Catalpa); anthers tetrasporangiate

or disporangiate. Tapetum amoe-

boid. Pollen grains 2-celled, in

monads, tetrads, or rarely polyads,

3(-4)-colpate, 3(-4)-colporate,

3-porate, zonocolpate, pericolpate

and syncolpate. Annular or some-

times cupular intrastaminal nectary

disc usually present. Gynoecium of

two carpels; style with 2-lobed

stigma; ovary superior, typically

2-locular with two placentas per

locule or sometimes 1-locular with

two or four more or less intruded

parietal placentas or rarely

(Tourrettia) 4-locular with numer-

ous and usually anatropous ovules

uniseriate in each locule. Ovules

numerous, ascending, anatropous,

hemitropous, or orthotropous.

Fruits septicidal or loculicidal,

bivalved capsules, very often with a

replum, or berries (in Crescentieae

and Coleae). Seeds in capsular

fruits usually ﬂ at, winged; wings

can be formed by fused capillary

hairs or usually by a thin mem-

brance (Hesse and Morawetz 1980);

seed coat exotestal (Corner 1976);

embryo straight, with usually

enlarged and more or less folia-

ceous cotyledons; endosperm

wanting. Present iridoid glycosides,

iridoid alkaloids, ﬂ avonols, and

ﬂ avones, and also ursolic and shi-

kimic acids, n = 11, 14, 15, 18, 20.

. . . . . . . . . . . . . . 7. bignoniaceae.

14 Seeds not ﬂ at and not winged.

Herbs, subshrubs, or shrubs,

rarely trees.

15 Style terminal or nearly so.

16 Calyx well developed.

17 Seeds with endo sperm. Perennial

or annual herbs, rarely shrubs or

subshrubs, with both simple and

multicellular and usually also

glandular hairs. Leaves decus-

sate, simple, entire to dentate or

rarely deeply pinnatiﬁ d, estipu-

late. Inﬂ orescence axillary of

one or more superposed cymes,

the latter 1-several-ﬂ owered or

ﬂ owers collected in spicate

bracteate inﬂ orescences; calyx

(4-)5-lobed, accrescent; corolla

sym petalous, slightly zygomor-

phic, unequ- ally 5-lobed, the

lobes imbricate; stamens didy-

namous, inserted low down in

the tube, included; anthers with

small apical connective, open-

ing longitudinally. Pollen grains

3-colpate. Ovary superior, 2-loc-

ular, ovule one per locule, anat-

ropous, axile at the base of the

ovary. Fruit indehiscent, oblong-

ovoid, compressed, shallowly

grooved between the two halves,

pubescent around the thickened

persistent base of the style; testa

of the seeds foveolate, the fove-

oles arranged in longitudinal

lines. . . . . . 27. nesogenaceae.

Superorder LAMIANAE 555

17 Seeds without endosperm. Large

scandent shrubs, sometimes stel-

late-tomentose. Young twigs

quadrangular or terete. Leaves

opposite, simple, entire or ser-

rate, or dentate, estipulate.

Flowers sessile, in 3–7(-9)-ﬂ ow-

ered capitate cymes, often aggre-

gate into large terminal or

axillary panicles, each cyme

with an involucre of more or

less membranous, colored, per-

sistent bracts and bracteoles,

bisexual, zygomorphic. Calyx

4–5–8-toothed, more or less

accrescent or inﬂ ated in fruit,

teeth open or scarcely valvate.

Corolla 6–16(-18)-lobed, lobes

subequal or bilabiate, imbricate.

Stamens 4–16(-18), alternating

with the corolla members. Pollen

grains 2-celled, 3-colpate, tec-

tate-perforate or rugulose, rarely

operculicolpate. Disc absent.

Gynoecium of two carpels, with

ﬁ liform style and biﬁ d stigma;

ovary superior, 2-locular (mor-

phologically), or 1-locular

(above); locules partially sec-

ondarily divided by ‘false septa’,

with two ovules pendulous from

the free-central placenta. Fruits

small, dry, indehiscent,

1–4-seeded drupe, more or less

included in the persistent calyx;

embryo straight. n = 12, 14, 17,

18. . . . 8. symphoremataceae.

18 Plants usually woody.

19 Stamens usually four (the ﬁ fth

one staminodial or absent),

inserted on the corolla, or

reduced to two. Trees or shrubs,

herbs, sometimes lianas, gla-

brous or pubescent, nonglandu-

lar trichomes usually unicellular.

Vessels with simple, or occa-

sionally scalariform perforation.

Nodes unilacunar, with 1-several

traces. Leaves opposite or rarely

verticillate or alternate, entire,

simple or pinnately or palmately

compound, lobed to dissected,

sometimes much reduced, scale-

like, or spinose, estipulate.

Stomata anomocytic or diacytic,

rarely paracytic. Flowers in vari-

ous types of racemose inﬂ ores-

cences, mainly cymose in the

Viticoideae, often with involucre

of colored bracts, mostly brac-

teolate, bisexual or rarely some

of them unisexual (dioecious).

Calyx 5(4)-lobed or toothed,

sometimes zygomorphic.

Corolla mostly 5-lobed, with

imbricate lobes, mostly more or

less zygomorphic, sometimes

bilabiate, often with narrow tube,

rarely campanulate. Anthers

2- locular, basiﬁ xed or dorsiﬁ xed.

Pollen grains with the thickening

of the exine immediately adja-

cent to the colpi or pores, 3-col-

pate or more often 3-colporate,

sometimes 4-colporate, rarely

3-pororate or 3-porate, tectate-

perforate and without supratectal

processes, psilate or irregularly

rough. Gynoecium of two or

rarely (Duranta) four carpels;

style with usually lobed stigma

and with prominent stigmatoid

tissue; ovary usually initially

2-locular but later commonly

subdivided by intrusion of parti-

tions from the carpellary midribs

into four uniovulate locelli;

rarely the anterior or superior

carpel suppressed. Ovules 1–2

per locule, erect or rarely pendu-

lous, mostly anatropous. Fruits

drupaceous with two or four

pyrenes or often of 1-seeded,

separating, nutlike mericarps.

Seeds with straight, oily embryo;

endosperm mostly wanting;

seed coat consists of several lay-

ers of thin-walled cells. Usually

556 Subclass VIII. LAMIIDAE

producing iridoid compounds

and often the phenolic glycoside

orobanchin, less often alkaloids,

contain ethereal oils, n = 5, 7,

10–12, 15–20, 22–26, 30.

. . . . . . . . . . . 24. verbenaceae.

19 Stamens 5–6, attached to the

corolla tube and alternating

with corolla lobes, unequal.

Evergreen small trees or shrubs

with two types of hairs: simple,

unicellular trichomes ﬁ lled with

calcium carbonate and accom-

panied by cystoliths in the adja-

cent epidermal cells at the base,

and bicellular, club-shaped glan-

dular hairs. Vessels with simple

perforations. Leaves opposite or

subverticillate, small, leathery,

simple entire, and sometimes

emarginate, estipulate. Stomata

anomocytic. Flowers axillary, in

1–7-ﬂ owered congested racemes

or dichasia, often reduced to

only 1–2 ﬂ owers, bisexual, but

functionally unisexual, dioe-

cious, slightly zygomorphic.

Calyx small, 5–6-partite, imbri-

cate or open, persistent. Corolla

5–6-lobed, tube broadly funnel-

shaped, lobes imbricate. Anthers

basiﬁ xed, introrse to extrorse, 2-

or 4-celled. Pollen grains 3-col-

porate, reticulate, with long

columellae and smooth colpus

membrane. Nectariferous disc

presents at the base of ovary in

female ﬂ owers. Gynoecium of

two united carpels; ovary supe-

rior, 1-locular, shortly stipitate,

with two parietal placentas, the

placentas bearing either one

basal erect ovule each, two basal

ovules each, or two basal ovules

on one and two subapical,

ascending or pendulous ovules

on the other. Fruits elongated,

many-ribbed, and sulcate,

1–4-seeded capsule, 2-valved

from the apex. Seeds long-lin-

ear, with a dense tuft of hairs at

the apex; embryo straight, lin-

ear, almost ﬁ lling the seed;

endosperm very thin, ﬂ eshy.

Colleters absent. Producing qui-

nol glycoside cornoside and two

compounds (lugrandoside and

echinacoside) belonging to ver-

bascoside group, but lack iri-

doids. 9. plocospermataceae.

18 Perennial, erect herbs from short,

vertical rhizome, non-aromatic,

with simple hairs. Young stems

tetragonal. Nodes unilacunar,

with three traces. Leaves oppo-

site, entire, coarsely dentate.

Stomata anomocytic, sometimes

anisocytic. Flowers in elongate,

narrow terminal and axillary

slender spikes, bracteate, small,

zygomorphic, bisexual, bracte-

ate, 5-merous. Calyx tubular,

5-toothed, the dorsal teeth

becoming spinose and hooked in

fruit, two ventral teeth

unchanged. Corolla tubular, bila-

biate, purple or blue. Stamens

four, inserted midway down the

corolla tube to in the throat of the

corolla tude, didynamous; ﬁ la-

ments glabrous; anthers subreni-

form. Pollen grains 3-celled,

tricolpate. Gynoecium of two

carpels, pseudo-monomerous

(the abaxial carpel is sup-

pressed); style with 2-lobed

stigma, without persistent stig-

matoid tissue; ovary oblique,

1-locular, with subbasal, erect,

orthotropous ovule; endosperm

haustoria chalazal. Fruits

1-seeded achenes enclosed in

the persistent accrescent calyx,

the testa adnate to the membra-

nous pericarp; seed coat vesti-

gial, embryo oblong, endosperm

scanty, consists of only two lay-

ers of cells, cotyledons convo-

Superorder LAMIANAE 557

lute. Iridoids detected, n = 7, 14.

. . . . . . . . . .25. phrymataceae.

16 Calyx absent or reduced to a

minute disc-like rim (in

Cyclocheilon physocalyx), its

function taken over by two

bracteoles. Shrubs or subshrubs

with red roots (?) and simple

and glandular hairs. Leaves

opposite or subopposite, simple,

entire, estipulate. Flowers soli-

tary on bracteate axillary pedi-

cels, bisexual; bracteoles

lateral, membranous, more or

less free or joined, ﬂ at or con-

vex and winged, their margins

closely adpressed and enclos-

ing the bud, accrescent.

Corolla obliquely funnel-

shaped, 5-lobed, lobes broad,

imbricate. Stamens four, didy-

namous; ﬁ laments long-pilose,

inserted on corolla tube; anthers

with a short connective. Pollen

grains 3-, rarely 4-colpate, tec-

tum very ﬁ nely perforate-rugu-

late. Gynoecium of two carpels;

style curved, with lingulate

stigma; ovary compressed,

completely or incompletely

2-locular or 1-locular, with

2–10 ovules on long funicles;

placentation axile or parietal.

Fruits very strongly com-

pressed discoid 1-locular cap-

sules with loculicidal

dehiscence (Cyclocheilon) or

breaking up into two 1-seeded

mericarps (Asepalum). Seeds

without endosperm. . . . . . . . .

26. cyclocheilaceae.

15 Style mostly gynobasic.

Perennial or annual herbs, sub-

shrubs, shrubs (sometimes eri-

coid), or less often trees, rarely

woody climbers, mostly aro-

matic. Stems erect to prostrate,

sometimes forming stolons or

large or slender rhizomes;

young stems and twigs usually

quadrangular. Indumentum of

glandular and non-glandular

trichomes, rarely scale-like,

dendroid or stellate. Leaves

opposite or sometimes verticil-

late, very rarely alternate, dis-

sected or entire, simple or

occasionally pinnately com-

pound, stipulate or estipulate.

Stomata mostly diacytic.

Inﬂ orescences of various types,

mostly composed of small,

compact, axillary cymes that

form a verticillaster at each

node, or sometimes the axillary

cymes reduced to single ﬂ ow-

ers; ﬂ owers mostly bracteolate,

bisexual or sometimes unisex-

ual, more or less strongly zygo-

morphic. Calyx 2- (e.g.

Prostanthera), or 3- (in

Melittis), or 4- (e.g. Preslia)

lobed, but basically 5-lobed or

dentate, tubular or more or less

bilabiate, seldom more or less

actinomorphic with four lobes,

as in Mentha (one lobe repre-

senting two connate lobes),

imbricate, persistent. Corolla

limb 5- or rarely 4-lobed,

mostly bilabiate, imbricate.

Stamens two or four, usually

didynamous, sometimes the

upper (posterior) pair stamino-

dial, rarely the lower pair ster-

ile (Mosla); ﬁ laments inserted

near the base of the corolla

tube, or midway down the

corolla tube, or in the throat of

the corolla tube, and sometimes

the pairs attached at different

levels; anthers 1–2-locular,

basiﬁ xed or dorsiﬁ xed, introrse,

opening longitudinally or

rarely by apical pores; the con-

nective often much developed.

Pollen grains 2- or 3-celled,

3(4)-colpate or 6(-12)-colpate,

558 Subclass VIII. LAMIIDAE

mostly suprareticulate, colpal

membrane mostly without

ectexine elements.

Intrastaminal nectary disc usu-

ally prominent, often devel-

oped on anterior side only.

Gynoecium of two carpels,

sometimes on gynophore; style

with 2-lobed stigma; ovary

4-lobed, fundamentally 2-locu-

lar, but early in development a

constriction appears in the

ovary in the antero-posterior

plane, longitudinally dividing

each carpel in half, so that the

ovary becomes divided into

four more or less distinct seg-

ments united by gynobasic

style. Ovules two per locule, or

solitary in each of four ovary

lobes, basal-axile, usually

anatropous to hemianatropous,

erect, often with funicular

obturator and usually with

endothelium; endosperm haus-

toria usually chalazal and

micropylar, the latter aggres-

sive. Fruits of (1-)4 one-seeded,

nutlike mericarps with a hard

pericarp or rarely mericarps

with a ﬂ eshy pericarp, rarely

druplets (Stenogyne,

Gomphostemma, Prasium).

Seeds with straight or rarely

bent (Catoferia, Scutellaria),

or slightly curved embryo; the

radicle directed downward;

endosperm wanting or scant,

ﬂ eshy endosperm often

absorbed by the developing

embryo. Characterized by very

broad spectrum of isoprenoid

compounds, caffeic acids

derivatives and apigenin- and

luteolin-derived ﬂ avonoids and

by the replacement of starch

accumulation in perennial parts

by oligo-saccharidcs; store

mainly oils rich in linolic or

linoleic acids in their starch-

free seeds; true tannins and

alkaloids are lacking; iridoids

present or absent, n = 5–11, 13,

15, 17, 19, rarely up to 120

(Salvia ombrophila – Harley

et al. 2004). . . 29. lamiaceae.

20 Plants characterized by muci-

lage trichomes with very

shortly stalked heads com-

posed of four or more cells, the

latter being convexly arched

outward from the apex of the

stalks, and the lamina almost

completely ﬁ lled with muci-

lage. Fruits very often with

hooks or horns or prickles, or

sometimes winged.

21 Plants terrestrial. Ovary

superior.

22 Flowers mostly with character-

istic glands (extraﬂ oral necta-

ries) representing abortive

ﬂ owers axillary to as many

bracts at the base of the pedi-

cel. Perennial or annular herbs,

rarely shrubs or small trees,

covered by mucilage-produc-

ing glandular hairs. Leaves

opposite, the upper ones alter-

nate, simple, entire, dentate or

lobed, estipulate. Stomata ano-

mocytic, anisocytic, or dicytic

and paracytic. Flowers solitary

or in terminal cymes (usually

3-ﬂ owered), bisexual, zygo-

morphic, 5-merous. Sepals

more or less united. Corolla

often with oblique limb, lobes

imbricate. Stamens four,

inserted near the base of the

corolla tube, and generally

paired, with small, subulate

staminodium in place of the

ﬁ fth (posterior) one; anthers

often connivent or contiguous

in pairs. Pollen grains

2-celled, 5–13 zonocolpate.

Intrastaminal, ﬂ eshy nectary

Superorder LAMIANAE 559

disc present. Gynoecium of

two carpels, with long style

and 2-lobed stigma; ovary

2–4-locular or apparently

1-locular, often with false

septa; ovules one (Josephinia),

or two to many in each locule,

pendulous, or horizontal, or

ascending, anatropous. Fruits

loculicidal carpels or nuts,

often with horns or hooks.

Seeds winged or not, testa mul-

tiplicative, exotestal cells pali-

sade or otherwise thickened,

mesotesta with crystals;

embryo straight, endosperm

scanty or absent. Contain iri-

doids, ﬂ avone glycoside, ver-

bascoisede-type compounds, n

= 8, 13, 16. .15. pedaliaceae.

22 Flowers without extraﬂ oral

nectaries. Perennial or annual

herbs, erect to prostrate, often

with tuberous roots. Arial parts

covered with glandular-viscid

trichomes. Leaves opposite or

alternate, petiolate, simple,

sticky, entire to sinuate or

almost lobed. Stomata anomo-

cytic or anisocytic, paracytic

and diacytic. Flowers in termi-

nal racemes, bisexual, zygo-

morphic. Sepals ﬁ ve, nearly

free or partly united, some-

times spathaceous. Corolla

5-lobed, lobes imbricate.

Fertile stamens four in two

pairs (the ﬁ fth reduced to a sta-

minodium) or less often

(Martynia) two (anterior ones);

anthers connivent, tetrasporan-

giate, opening longitudinally.

Pollen grains not colpate, retic-

ulate (Martynia), areolate

(Proboscidea and Ibicella),

and “ringlike” (Craniolaria);

exine dissected into 20–40

platelets. Intrastaminal nectary

disc glandular. Gynoecium of

two carpels, with long style and

2-lobed stigma; ovary 1-locular

with two massive parietal

T-shaped placentas divided into

two divaricate lobes ovules few

to numerous, anatropous. Fruits

usually hooked at the end or

horned, loculicidal after the

separation of a soft, deciduous

exocarp, mesocarp more or

less ﬂ eshy, falling off, endo-

carp wood. Seeds with straight

embryo and scanty endosperm.

Contain iridoids, flavone

glycosides and verbascoside-

type compounds, n = 15, 16.

. . . . . . . . . 16. martyniaceae.

21 Plants aquatic. Ovary inferior.

Perennial herbs with creeping

rhizomes and more or less

ﬂ oating, elongate slender

stems. Leaves opposite, dimor-

phic, the submerged ones

oblong, remotely serrate, the

upper (ﬂ oating) deltoid-rotun-

date or reniform-cordate,

crenate. Both side of the leaves

covered by glandular hairs.

Stomata anomocytic, exclu-

sively occur on the upper face

of the ﬂ oating leaves. Flowers

solitary, axillary, on slender

pedicels, bisexual, zygomor-

phic, chasmogamous from

ﬂ oating leaves, cleistogamous

from submerged leaves. Calyx

5-lobed, imbricate, persistent,

with ﬁ ve hornlike appendages

near the apex, these elongate

after anthesis. Corolla tubular,

5-lobed, slightly bilabiate,

imbricate. Two posterior sta-

mens fertile, the other two sta-

minodial; ﬁ laments ﬁ liform;

anthers on large peltate con-

nective, included. Pollen grains

3-zonocolpate. Gynoecium of

two carpels, with slender style

and dilated, shortly 2-lobed

560 Subclass VIII. LAMIIDAE

stigma; ovary 2-locular, poste-

rior locule fertile, with two

pendulous, anatropous ovules,

the anterior one abortive and

empty. Fruits narrow-elongate,

1-seeded, indehiscent, crowned

with ﬁ ve spreading, rigid

appendages below the calyx,

subulate and spinose. Seeds

linear, 4-ridged; seed coat exot-

estal (Singh 1960); embryo

straight, endosperm scanty. n =

ca. 25. . . . 17. Trapellaceae.

20 Plants without mucilage

trichomes.

23 Plants characterized by scat-

tered secretory cavities lined

with epithelium, which occur

in the leaves and in the primary

cortex and/ or pith of the axis.

Shrubs or sometimes small

trees, often with lepidote or

plumose indumentum or with

gland-tipped hairs. Nodes uni-

lacunar, internal phloem

absent; primary vascular tis-

sue centrifugal. Leaves alter-

nate or seldom opposite, entire

or dentate, often more or less

leathery, petiolate to sessile,

often (when secretory cavities

large) with pellucid or raised

dots. Flowers in small axillary

cymes or solitary in the axils,

bisexual. Sepals ﬁ ve, connate,

lobes imbricate or open.

Corolla from essentially acti-

nomorphic (as in Myoporum)

to strongly zygomorphic (as in

Bontia), often bilabiate, lobes

imbricate. Stamens mostly

four (the ﬁ fth posterior repre-

sented by a staminodium or

absent) or rarely ﬁ ve. Anthers

connivent or separate, introrse,

conﬂ uent at apex, opening

with longitudinal slits or trans-

versely. Pollen grains 2–4-col-

porate, often with two pores in

each colpus. Gynoecium of

two carpels, with terminal,

sometimes impressed style

and simple stigma; ovary supe-

rior, 2-locular, rarely 4-locu-

lar, with (1)2 anatropous

ovules pendulous near the

summit of the partition in each

locule, or 4–8 ovules per loc-

ule, superposed in pairs, or the

locules subdivided into 4–10

uniovulate compartments.

Fruits indehiscent drupaceous

or separating into 1-seeded,

drupe-like segments, the

exocarp dry and papery and

separable from the thin carti-

laginous or thickened endo-

carp by mesocarp (Henrickson

and Flyr 1985). Seeds

unwinged or rarely (Androya)

winged, with more or less

straight embryo; endosperm

scanty or lacking. Contain iridoid

and verbascosides, n = 17, 18.

. . . . . . . . .18. myoporaceae.

23 Plants without scattered secre-

tory cells. Lamina without

secretory cavities. Internal

phloem present, or absent.

24 Flowers ebracteate and mostly

without bracteoles. Perennial

suffrutescent herbs or shrubs,

decumbent to ascending, or

erect, glabrous or more often

leaves and branchlets provided

with short, glandular, and long,

multicellular and usually nong-

landular hairs. Internal phloem

present; primary vascular tis-

sue bicollateral. Leaves oppo-

site, sometimes alternate above,

simple, lanceolate to ovoid,

acuminate, dentate to serrate,

not gland-dotted. Stomata ano-

mocytic. Flowers in the axil of

the upper leaves, subsessile or

with short pedicels, solitary,

bisexual, zygomorphic. Calyx

Superorder LAMIANAE 561

deeply and almost regularly

5-lobed, persistent, lobes are

covered with glandular hairs.

Corolla with rather narrow

cylindrical tube and ﬁ ve lobes

that overlap in bud; corolla

tube glabrous or often shortly

glandular and puberulous on

the outer side, and closely cov-

ered with hairs pointing for-

ward on the upper half of the

inner side. Stamens four

inserted on the corolla tube;

ﬁ laments short and slender;

anthers comparatively large.

Pollen grains 4-colporate,

foveolate, with small perfora-

tions. Gynoecium of two car-

pels; style short, with the

stigma directed upward; ovary

2-locular, with 4(-6) ovules per

locule, pendulous from the

upper part of the septum. Fruits

2-locular (or by abortion 1-loc-

ular) globose drupes with one

pyrene and extremely hard,

thick inner walls. Seeds with

stout, clavate funicle and hard,

black testa; endosperm want-

ing, n = 19. . . .19. oftiaceae.

24 Flowers bracteate and

bracteolate.

25 Ovules anatropous to campy-

lotropous, each with a more or

less strongly modiﬁ ed funicu-

lus. Mostly perennial herbs,

subshrubs, and shrubs, rarely

trees, very diverse in habit,

including lianas, xerophytes,

and aquatics (even mangrove

Acanthus ilicifolius). Provided

with various types of trichomes.

Internal phloem present, or

absent. Leaves usually well

developed or much reduced,

opposite or (Nelsonioideae)

alternate, entire or pinnately

lobed, gland-dotted or not gland-

dotted, simple. Cystoliths very

commonly present, showing as

streaks in the lamina. Stomata

always almost diacytic. Flowers

in various kinds of cymose or

less often racemose inﬂ ores-

cences or sometimes solitary,

the bracts and bracteoles often

showy, bisexual. Calyx more or

less deeply (4)5–16-lobed, or

lobes sometimes much reduced

(as in Thunbergia), lobes imbri-

cate or valvate, or contorted, or

open in bud. Corolla more or

less zygomorphic, usually typi-

cally 5-lobed and usually bila-

biate (upper lip sometimes not

developed, as in Acanthus),

lobes imbricate or contorted.

Stamens usually four or two

and paired (upper stamens rep-

resented by staminodia), rarely

all ﬁ ve stamens fertile

(Pentstemona-canthus). Anthers

separate from one another, or

connivent, dorsiﬁ xed, often with

one lobe reduced or abortive,

unilocular to bilocular. Pollen

monosiphonous, shed as single

grains. Pollen grains 3-colpate,

or porate, or colporate, or foram-

inate; 2-celled, or 3-celled.

Annular or glandular intrastami-

nal disc usually present.

Gynoecium of two carpels, with

slender style and 2-lobed and

variously shaped (often funnel-

form) stigma; ovary superior,

usually 2-locular, with two or

more (in Nelsonioideae more or

less numerous) ovules per loc-

ule on each intruded parietal

placenta. Fruits usually loculi-

cidal, explosive capsules or

(Mendoncieae) drupes. Seeds

with large, straight, or more or

less curved embryo and usually

without endosperm, but in

Nelsonioideae endosperm more

or less well developed, oily, and

562 Subclass VIII. LAMIIDAE

ruminate; in many genera seeds

with often hook-like funicular

projection (retinaculum or jacu-

lator). Iridoids detected, alka-

loids present or absent; ﬂ avonols,

when present, kaempferol and

quercetin; n = 7–21.. . . . . . . . . .

20. acanthaceae.

25 Ovules orthotropous to hemi-

anatropous. Trees or shrubs of

tropical coasts with pneumato-

phores. Branchlets and twigs

with jointed appearance from

swollen nodes. Secondary thick-

ening anomalous. Intraxylary

phloem present. Axial paren-

chyma apotracheal. Nodes trila-

cunar with three traces. Leaves

opposite (decussate), entire,

coriaceous, leathery; simple,

estipulate, with salt glands on

both sides. Stomata diacytic.

Flowers sessile, in terminal or

axillary pubescent spicate or

subcapitate inﬂ orescences,

subtended by a single bracts

and a pair of bracteoles, small,

bisexual, actinomorphic. Calyx

cup-shaped, deeply 5-lobed,

lobes imbricate. Corolla acti-

nomorphic or variously zygo-

morphic, campanulate-rotate,

4(5–6)-lobed, rotate or campan-

ulate, yellow, lobes imbricate.

Nectary disc inconspicuous.

Stamens mostly four, equal or

subdidynamous, inserted in the

throat of the corolla tube;

anthers sessile. Pollen grains

3-colporate, tectate-perforate

to microreticulate, psilate, col-

pal membrane granular.

Gynoecium of two carpels;

style with biﬁ d stigma; ovary

superior, 2-locular; locules

partially secondarily divided

by false septa, with a free-cen-

tral more or less winged pla-

centa; ovules two per locule

(one per locellus), hanging

from tip of central placenta,

orthotropous to hemianatro-

pous, unitegmic, but with

incomplete growth of the integ-

ument, which does not form a

micropyle; endosperm hausto-

ria chalazal and micropylar.

Endothelium wanting. Fruits

compressed capsule dehiscent

by two thick, leathery valves,

by abortion regularly 1-seeded,

bracteoles and sepals persis-

tent. Seeds imperfectly covered

by its testa, with viviparous,

green embryo; cotyledons

folded around the plumular

axis, radicle fully or partly

hairy, endosperm lacking.

Contain iridoid glycosides, n =

18, 32. . . .21. avicenniaceae.

7 Plants insectivorous.

26 Flowers actinomorphic, nectary

spur absent. Evergreen shrubs,

subshrubs or perennial insectiv-

orous herbs with woody rhizome

bearing in any one season the

dying parts of the previous and

the growing ones of the present

season. Trapping mechanism

passive. Indumentum consists of

numerous glands of two types –

sessile and stalked; the stalked

glands, which are found even

on the ovary wall, have a muci-

laginous head of usually 32

cells radiating from the center

and standing out like an

umbrella top; the apex of the

stalked head traps small insects,

while the sessile glands digest

them. The glands closely

resemble those of Pinguicula

in the Lentibulariaceae (Conran

and Carolin 2004). Vessels

mostly with simple perforations,

but scalariform perforations

with 1–3 bars are also found;

Byblis gigantean has unusual,

Superorder LAMIANAE 563

narrow, tracheid-like vessels;

lateral pitting alternate. Fibers

consisting of true tracheids.

Rays mostly biseriate or triseri-

ate, composed of erect cells and

very few are square. Axial

parenchyma sparse, diffuse.

Sieve-element plastids of

Ss-type. Nodes unilacunar with

one or three traces. Leaves alter-

nate, elongate linear, terete or

ﬁ liform, crowded, circinate in

vernation, estipulate. Stomata

paracytic. Flowers solitary in

the axils of leaves, bisexual,

ebracteolate, 5-merous (except

gynoecium). Sepals imbricate,

basally connate, persistent.

Petals imbricate or contorted,

basally shortly connate, broad-

cuneate, apically ﬁ mbriate, or

entire (B. gigantea). Stamens

ﬁ ve, alternipetalous, inserted at

the base of petals or free, often

declinate; ﬁ laments short, subu-

late; anthers basiﬁ xed, tetraspo-

rangiate, introrse, bilocular,

opening by apical pore or very

short, porelike slit, with a glan-

dular binucleate tapetum. Pollen

grains 2-celled, 3–4- colpate,

3-colporate, or 4-, or 6-rugate.

Gynoecium of two united car-

pels, with elongate, ﬁ liform

style and capitate or oblong

stigma; ovary superior and as a

result of the fusion of two pari-

etal placentas 2-locular, with

10–50 ovules per locule. Ovules

pendulous, anatropous, uniteg-

mic, tenuinucellate, with

endothelium. Endosperm cellu-

lar, with both chalazal and

micropylar haustoria. Fruits

2-locular, 2–4-valved, loculi-

cidal, smooth capsules with

many seeds. Seeds small, with

coarsely verrucose testa,

straight, elongate, and thin, lin-

ear embryo with small but

differentiated cotyledons, sur-

rounded by copious, starchy

endosperm, occasionally with

deposits of calcium oxalate;

seed coat consists of 2–3 lay-

ers of cells of pachytesta.

Iridoids present, but iridoid

type is unknown, n = 7, 8,

. . . . . . . . . 9. 22. byblidaceae.

26 Flowers zygomorphic, nectary

spur present. Insectivorous herbs

found in water or other wet habi-

tats, occasionally epiphytic,

rooted in the substrate or rootless

and free-ﬂ oating with the photo-

synthetic organs submerged.

Trapping mechanism active or

passive. Provided with stalked

and/or sessile glands. Stem in

Utricularia bearing bladders

with a trap mechanism capturing

small animals. Nodes unilacu-

nar. Leaves alternate, linear to

broadly ovate, entire, estipulate,

in basal rosettes in Pinguicula

and Genlisea, which has also

tubular trap-leaves arising sepa-

rately from the rhizome. Stomata

diacytic or rarely anomocytic or

anisocytic, sometimes absent.

Flowers in a bracteate raceme

(rarely 1-ﬂ owered) or solitary on

a bractless scape (Pinguicula),

bisexual. Calyx equally 4–5-

lobed or more or less 2-cleft,

persistent, imbricate or open in

bud. Corolla bilabiate or more or

less 5-lobed, lobes imbricate, the

lower lip basally more or less

spurred or saccate. Stamens two

(anterior pair), inserted near the

base of the corolla tube; anthers

connivent, dorsiﬁ xed, unilocular

or 2-locular. Pollen grains

3–4-colporate or multicolporate,

in tetrads or in spiroapertutate

monads. Nectary disc absent.

Gynoecium of two carpels with