Takhtajan Armen. Flowering Plants

Подождите немного. Документ загружается.

262 Subclass V. DILLENIIDAE

to many carpels; style with a

lobed or capitate stigma; ovary

superior, plurilocular or

(Goethalsia, Mollia) unilocular,

with (1)2-several ovules on axile

or intruded parietal placentas in

each locule. Fruits dry or ﬂ eshy,

dehiscent or not. Embryo mostly

straight, n = 7–41. 2. tiliaceae.

8 Endosperm scanty or none.

9 Stem with characteristic verti-

cal branching system of inter-

cellular resin canals in the pith,

wood, and bark. Ectomycorrhizal

trees. Secretory cavities in pith

absent. Rays heterogeneous to

homogeneous, larger rays

multiseriate. Flowers without

androgynophore. Sepals val-

vate, often two or more of them

conspicuously accrescent and

wing-like in fruit. Anthers bas-

iﬁ xed. Pollen grains 3-colpate,

the exine convoluted, without

clear columellae. Ovary

(2)3-locular, each locule with

two ovules; micropyle endos-

tomal. Produced terpenoids,

sesquiterpene oleoresins and

dipterocarpol. n = 6, 7, 10, 11(-

13). . . . 3. dipterocarpa ceae.

9 Stem without resin canals.

Secretory cavities in pith pres-

ent. Rays heterogeneous, uni-

seriate or sometimes biseriate.

Flowers with or without

androgynophore. Sepals imbri-

cate or less often valvate, equally

accrescent, sometimes winglike.

Anthers basiversatile. Pollen

grains 3-colporate, the exine

tectate- columellate. Ovary

2–5-locular, each locule with

1–2 or 4 ovules. Fruits 1-seeded,

nutlike, or 3–5 locular capsules.

Produced sesqui- and triterpe-

noids.. . . . . . . .4. monotaceae.

7 Pollen grains in tetrads, 3–6-colpate.

Stamens usually very numerous,

rarely (spp. of Leptolaena) 5–10,

usually persistent, generally inserted

inside an entire or toothed, more or

less cupular nectary disc of stamino-

dial origin; ﬁ laments free or weakly

connate at the base into 5–10 fasci-

cles; anthers basi- or dorsiﬁ xed,

opening longitudinally. Gynoecium

of 3–4(5) carpels with a terminal

style and expanded, usually lobed

stigma; ovary with as many locules

as carpels, with (1)2-many ovules

per carpel; ovules apical-descendent,

basal-ascendant, or axile in the mid-

dle of the ovary, anatropous. Petals

contorted, 5(-6), large, free or slightly

united at the base. Sepals mostly

3(4–5), imbricate-contorted, when

more than three, the outer ones

smaller than the inner three. Flowers

in umbel-like or more or less panicu-

liform inﬂ orescence or rarely soli-

tary, bisexual, usually subtended by

an involucel (epicalyx) of bracteoles

more or less connate to form a cup;

in Eremolaena epicalyx lacking.

Fruits loculicidal capsules or inde-

hiscent and with only one or a few

seeds. Seeds hairy or not, with

straight embryo; seed coat exoteg-

mic; endosperm usually copious and

starchy, sometimes ruminate or

reduced; cotyledons cordate. Leaves

simple, entire, usually persistent,

with usually caducous stipules.

Nodes trilacunar. Trees or shrubs,

often with stellate hairs. Presence of

sapinins, glucosides of myrcetin

and other ﬂ avonols, phenolic acids

of the cinnamic and the benzoic

series, n = 11. 5. sarcolaenaceae.

6 Mucilage cells and cavities absent.

Phloem not stratiﬁ ed.

10 Trichomes glandular with globular

heads. Small to medium sized trees

or shrubs; pubescent of stellate,

long unbranched, and glandular

hairs, especially on young growth.

Perforation plates non-bordered or

Superorder MALVANAE 263

minimally bordered. Axial paren-

chyma diffuse, scanty vasicentric,

diffuse-in-aggregated. Leaves

alternate, distichous, petiolate,

blades asymmetrically cordate at

base, palminerved, their margin

serrate; stipule-like appendages

ﬁ liform or foliaceous and peltate.

Flowers in supra-axillary position,

solitary of in few-ﬂ owered clusters,

pedicellate, actinomorphic, usually

bisexual. Sepals (4-)5(-7), valvate

in bud, with more or less spreading

tips, basally fused, forming a sau-

cer- to cup-like tube. Petals (4)5(-

7), more or less crumpled in bud,

imbricate, free, longer than the

calyx, thin, caducous, the outer

margin irregular. Stamens numer-

ous; ﬁ laments ﬁ liform, free or

almost so; anthers dithecal, some-

times versatile, ﬁ xed at or near the

base of the connective, dehiscing

by longitudinal slits, sometimes

restricted to their apical portion.

Pollen grains 3-colporate. Ovary

superior to inferior, syncarpous,

5-many locular, sometimes 1-locu-

lar in the upper part, lobed, pendu-

lous; style thick, sometimes almost

lacking; stigma thick, lobed-sulcate

to decurrent. Ovules numerous.

Fruits indehiscent, baccate. Seeds

numerous, small; embryo short,

straight. Endosperm composed of

large, thin-walled cells, oily.

Contain gallic and ellagic acids,

n = 14, 15. . . . 1. muntingiaceae.

10 Trichomes stellate or simple, not

glandular. Prostrate, tomentose,

perennial and annual herbs with

stellate trichomes; stems sympo-

dial, more or less woody below.

Vessels with simple perforations;

lateral pitting opposite, sometimes

nearly alternate. Have lysigenous

mucilage ducts in the pith. Sieve-

element plastids of Pcs-type.

Leaves alternate, simple, toothed,

lobed to pinnatiﬁ d; estipulate.

Stomata anomocytic. Flowers soli-

tary on axillary peduncles, bisex-

ual, actinomorphic, sometimes

with an epicalyx of ﬁ ve bra cteoles,

5-merous. Sepals and petals united

into a ﬂ oral tube that is dry at matu-

rity. Calyx lobes more or less val-

vate, corolla lobes imbricate or

contorted. Stamens 10, borne on

the ﬂ oral tube; ﬁ laments elongate,

subulate, glabrous or rarely pubes-

cent, more or less persistent;

anthers small, introrse, dorsiﬁ xed,

opening longitudinally. Pollen

grains 3-celled, 3-colporate, of a

unique bipolar type, with 3(4)

pores at each end. Gynoecium of

10 or (Grielum) 5–10 united car-

pels, with free subulate or ﬁ liform

stylodia that eventually become

indurated; stigmas small, capitate;

ovary more or less inferior, mul-

tilocular, but 2–4 of the locules on

the side toward the peduncle more

or less reduced, or their ovules not

maturing, with one ovule in each of

the fully developed locules. Ovules

pendulous, anatropous, apotropous.

Fruits dry, laterally membranous-

winged or spinose-muricate, car-

pels dehiscing ventrally, stylodia

sometimes spinescent. Seeds six to

several; exotegmic cells tangen-

tially elongated, crystalliferous,

other tegmic cells persistent;

embryo contains protein and oil,

but lacks starch; endosperm lack-

ing. Cyclopropenoid fatty acids

present; n = 7. . .6. neuradaceae.

5 Stamens generally connate into a tube

around the ovary, often seated on an

androgynophore, basically in two cycles,

the outer usually staminodial or some-

times suppressed, the inner often

branched. Anthers opening by longitudi-

nal slits or rarely by apical pores. Pollen

grains from 3-colporate and smooth or

reticulate to periporate and spinulose.

264 Subclass V. DILLENIIDAE

Gynoecium usually of ﬁ ve carpels, with

stylodia free or more or less connate into

a style. Ovary with as many locules as

carpels or rarely unilocular through the

failure of the parietal placentas to meet in

the center; in the tribe Sterculieae the

carpels united only by their stylodia and

becoming free at maturity, or wholly dis-

tinct from the beginning (as in Cola), but

this kind of apocarpy is most probably of

secondary origin (Takhtajan 1966).

Ovules (1)2 to many per locule, free car-

pel or free placenta. Fruits of various

types. Seeds sometimes arillate, with

straight or curved embryo and usually

copious endosperm or rarely (as in Cola)

without endosperm. Flowers bisexual or

less often unisexual, usually actinomor-

phic. Sepals valvate. Petals often small

and reduced, contorted, sometimes

absent, as in Sterculieae. Leaves alter-

nate, from simple and entire to palmately

lobed and palmately compound, stipu-

late. Stomata mostly anomocytic. Trees,

shrubs (sometimes lianoid), or rarely

herbs. Vessels with simple perforation of

rarely (Lasiopetalum) with rudimentarily

scalariform, n = 15, 16, 18, 20, 21 +.

. . . . . . . . . . . . . . . . .7. sterculiaceae.

4 Style more or less gynobasic.

11 Sepals and petals 5(6). Stipules broad

and coiled around the terminal bud,

caducous, and leaving a horizontal scare.

Leaves simple, entire, pinnately veined,

densely minutely pellucid-punctate,

smelling of camphor when crushed.

Stomata cyclocytic. Flowers in terminal,

few-ﬂ owered, paniculiform inﬂ ores-

cences, large, bisexual, fragrant. Sepals

free, widely imbricate, unequal (the

outer smaller), persistent until the begin-

ning of fruit development. Petals free,

large, slightly unequal, widely imbricate,

caducous. Stamens very numerous (up to

435 – Capuron 1963), free; ﬁ laments ﬁ li-

form; anthers basiﬁ xed, opening later-

ally by longitudinal slits. Pollen grains

small, spheroidal, 3- colporate.

Gynoecium of 2–4 free carpels with

common central elongate gynobasic

style with punctiform stigma; carpels

ovoid, very verrucose and with numer-

ous small, peltate glands. Ovules two in

each carpel, basal, ascending, with

micropyle facing outward. Fruits of 1–3

free 1-seeded mericarps with thin, cori-

aceous pericarps and with many small

glands; seed coat thin, no palisade layer,

inwardly-curving exo tegmen or

hypostase plug; embryo with thick cot-

yledons; endosperm absent. Shrubs or

small trees. . . . . . 8. diegodendraceae.

11 Sepals and petals four. Stipules large,

caducous. Leaves simple, entire, pubes-

cent or glabrous, pinnately veined or

3-veined, not pellucid-punctate and not

smelling of camphor when crushed.

Stomata anomocytic or anisocytic.

Flowers in axillary or terminal cymose

inﬂ orescences, bisexual. Sepals, unequal,

much imbricate. Petals unequal, imbri-

cate, densely streaked with short, resin-

ous lines, caducous. Stamens numerous

(more than 100), in 2–4 series, free or

inserted below a large, thick, gynophore-

like, cupular, wrinkled, denticulate

nectary disc; ﬁ laments slender, elongate,

more or less connate at the base, resin-

ducted; anthers small, introrse, loculi

short and widely separated by a broad,

glandular connective, opening longitudi-

nally. Pollen grains more or less spinose,

3–5-, rarely 6-colpate. Gynoecium of

two or four, or rarely three or ﬁ ve car-

pels; ovary superior, but partly immersed

in the nectary disc, usually 2-locular and

vertically more or less deeply lobed (in

Dialyceras resulting separation of car-

pels) with simple style inserted between

the lobes; stigma more or less capitate.

Ovules usually three in each locule, sub-

basal, anatropous, epitropous and

ascending. Fruits more or less deeply

lobed or globose, densely muricate, in

Dialyceras divided into mericarps. Seeds

large, with straight embryo; endosperm

ruminate or not, copious, oily, contains

elliptic starch grains (Bayer 2003);

Superorder MALVANAE 265

cotyledons cordate and bilobed apically.

Trees or shrubs. Leaves and seeds of

Rhopalocarpus) contain saponins, n =

19 (Rhopalocarpus lucidus – Sarkar

et al. 1982). . . 9. sphaerosepalaceae.

3 Anthers opening by pores or pore-like slits.

12 Leaves simple and entire, long-petiolate,

stipulate. Flowers in terminal thyrsoids,

large, actinomorphic. Sepals ﬁ ve, free,

imbricate, caduceus. Petals ﬁ ve, free,

imbricate, alternating with sepals.

Stamens numerous, free; anthers dithe-

cal, horseshoe-shaped opening with

short apical slits. Gynoecium of two,

rarely three carpels; ovary superior;

ovules many per carpel, anatropous,

pseudo-crassinucellate; micropyle zig-

zag. Fruits 2–4-valved setose capsules.

Seeds more or less straight, with a vesti-

gial funicular aril; endosperm starchy;

testa pulpy, eventually drying up and

minutely papillate from the bixin cells.

Small trees or shrubs, The leaves con-

tains several sesquiterpenes, elagic acid

and ﬂ avonoids, n = 7.. . . .12. bixaceae.

12 Leaves palmatisect, palmately lobed or

subentire- reniform, glabrous or pubes-

cent. Inﬂ orescence terminal or very rarely

axillary, thyrsoid. Anthers with one or

two apical and sometimes also with two

small basal pores. Gynoecium of 3–5 car-

pels; ovules many in each carpel; micro-

pyle zig-zag. Fruits 3–5-valved capsules,

the inner and outer layers separating and

forming alternative valves. Seeds large,

usually cochleate-reniform, without an

aril; endosperm oily; testa densely pilose

with white hairs along the curved dorsal

side of the seed. Seeds of Cochlospermum

possess cotton-like whitish or reddish

hairs, whereas those of Amoreuxia gla-

brous, pilose or echinate. Trees, shrubs, or

occasionally rhizomatous perennial herbs

with woody subterranean trunks. Present

ellagic acid, n = 6 (Cochlospermum)

. . . . . . . . . . . . . 13. cochlosperma ceae.

2 Nodes unilacunar.

13 Flowers with extrastaminal disc. Trees or

large shrubs (Tepuianthus auyantepuien-

sis) with resinous cells, and usually

densely pubescent with simple thrichomes.

Vessels with simple perforations. Fibers

with distinct bordered pits. Rays homoge-

neous, mostly uniseriate. Axial paren-

chyma scanty, vasicentric. Leaves

alternate or opposite, small, simple, entire,

pinnately veined, estipulate. Flowers

small, in cymose inﬂ orescences (terminal

or in the uppermost axils), androdioecious

or rarely bisexual, actinomorphic, recep-

tacle thickened. Sepals ﬁ ve, free, imbri-

cate, densely sericeous abaxially, glabrous

adaxially. Petals ﬁ ve, free, imbricate, tend

to be clawed, glabrous, yellowish. Nectary

disc consisting of 5–10 contiguous, sub-

orbicular, ﬂ eshy glands. Stamens 12–16

or 5, in 1–3 cycles, when unicyclic alter-

nipetalous; anthers 2-locular, sagittate,

versatile, introrse, dorsiﬁ xed, with more

or less appendaged connective. Pollen

grains 3–6-colporate, with prominent

ornamentation. Gynoecium of three

united carpels with free, forked stylodia;

ovary superior, 3-locular, with one pendu-

lous ovule per locule; placentation apical.

Ovules anatropous with ventral raphe, the

micropyle directed upward and outward.

Fruits densely sericeous, 3-locular, bony,

loculicidal capsules. Seeds with conspic-

uously ridged raphe, small embryo and

copious endosperm; the cotyledons poorly

differentiated (Boom and Stevenson

2004). . . . . . . . . . 15. tepuianthaceae.

13 Flowers without extrastaminal disc.

14 Pollen grains 3-colporate. Plants with

glandular, or simple, clustered, or

stellate hairs and colorless juice.

Nodes with one trace. Leaves opposite

or less often alternate or verticillate,

often more or less reduced, simple,

entire, petiolate to sessile, stipulate

or estipulate, often possess cystoliths.

Phloem not stratiﬁ ed. Axial paren-

chyma absent or very scanty. Flowers

solitary or in cymose inﬂ orescences;

sepals 3 or (4)5, when ﬁ ve the inner

contorted, often persistent. Anthers

basiﬁ xed. Gynoecium of 5 (6–10)

266 Subclass V. DILLENIIDAE

carpels in Cistus, in all other genera of

three carpels. Ovules mostly orthotro-

pous, rarely hemitropous or anatro-

pous. Seeds very small; seed coat of

two integuments, the outer thin, the

inner very hard; embryo usually curved

or bent into a hook or ring, or folded or

circinately more or less coiled, rarely

almost straight; endosperm starchy.

Shrubs, subshrubs, or herbs. Contain

glycosides of common ﬂ avonoids,

including myricetin and other ﬂ a-

vonols, and condensed and hydrolys-

able tannins, n = 5, 7, 9–12, 16, 18, 20,

24.. . . . . . . . . . . . . . . . 14. cista ceae.

14 Pollen grains mostly 8–60-porate.

Trees, shrubs, lianas, or rarely herbs,

evergreen or deciduous, usually poi-

sonous; wood often ﬂ uoresces. Stems

usually with intraxylary phloem

(except in the some ericoid or mosslike

species of Drapetes, and the anoma-

lous Lethedon and Solmsia) and in nine

genera (including Aquilaria) with

interxylary (included) phloem of

“foraminate” type. Leaves alternate or

opposite, not glandular-punctate, peti-

olate or sessile, simple, entire, estipu-

late. Stomata cyclocytic, rarely

(Octolepis) anomocytic. Flowers

bisexual or polygamous or dioecious,

actinomorphic, but only some species

of Lachnaea zygomorphic. Calyx 4–5-

lobed, tube campanulate or cylindric;

lobes imbricate to valvate. Corolla rep-

resented by free or united petaloid

appendages, isomerous and alternating

with the calyx lobes, less often double

or treble in number, entire or lobed,

rarely united into a ring, inserted at the

throat of calyx tube or at the middle or

near the base, or corolla absent.

Stamens equal or twice as many as the

calyx lobes or very rarely reduced to

two or even one, inserted below the

petals; in two or in one cycle, if in two

cycles then at two different levels (the

upper cycle antesepalous); ﬁ laments

ﬁ liform or slightly ﬂ attened, entirely or

partly adnate to the calyx tube; anthers

basiﬁ xed or rarely dorsiﬁ xed, introrse

or rarely extrorse. Pollen grains with

more or less clearly expressed “croto-

noid” ornamentation or rarely

(Octolepis) 3–6-porate with reticulate

ornamentation, somewhat resembling

those of some Tiliaceae (Erdtman

1952; Archangelsky 1966, 1971).

Nectary disc mostly present, annular,

cupular, membranous, or of 4–5 scales.

Gynoecium of (2)3–5, rarely 6–8, or

10–12 carpels (Lethedon), 3–5 carpels

(Solmsia, Deltaria, and Octolepis), or

more often of two carpels; style elon-

gate or short, sometimes very short, not

contorted, terminal or sublateral.

Stigma usually capitate, sometimes

papillose. Ovary sessile or shortly stip-

itate, 1–2-locular (rarely 3–12-locular),

with one pendulous ovule per locule.

Fruits loculicidal capsules or more

often indehiscent, nut-like, baccate, or

drupaceous, usually enclosed in the

base of the persistent calyx tube. Seeds

usually with a caruncle-like or tail-like

appendage, without an aril; exotegmen

and endotegmen with brown contents,

reticulately thickened and ligniﬁ ed;

embryo straight, white; endosperm

scanty or absent, rarely copious;

cotyledons large. Thymelaeoidea

contain highly toxic diterpene esters,

which often occur in the whole

plants, n = 7(-9)-10 (Thymelaeoideae).

. . . . . . . . . . . . 16. thymelaea ceae.

1 Anthers disporangiate and 1-locular.

15 Pollen grains generally smooth or merely

rugose, only rarely minutely spinose. Fruits

loculicidal capsules, rarely ﬂ eshy or inde-

hiscent. Trees, sometimes spiny, buttressed

or with swollen trunk, often with verticil-

late branches, or rarely shrubs or epiphytic

stranglers. Indumentum of stellate, excep-

tionally lepidote trichomes, or absent.

Leaves simple, rarely lobed, or digitate,

stipulate. Flowers actinomorphic or rarely

more or less zygomorphic, solitary of in

axillary clusters, rarely in cymes or paniculate,

Superorder MALVANAE 267

or terminal or leaf-opposed. Epicalyx of

3(2) distinct bracts usually present; sepals

commonly ﬁ ve, usually fused to more than

half their length, forming copular to tubular

calyx, sometimes persistent or even

accrescent, exceptionally imbricate in bud;

petals usually contorted, generally fused at

base with staminal column, often many

times longer than sepals and exceeding

calyx before anthesis; stamens usually

numerous (up to more than 1,000) or some-

times only ﬁ ve, generally forming a stami-

nal tube with antepetalous (or more or less

laterally shifted) lobes or phalanges and/or

distally free ﬁ lament portions; anthers usu-

ally monothecal, sometimes septate, straight

or reniform or spirally contorted; stamin-

odes absent or staminal tube exceptionally

with sterile appendages or collar-like thick-

ening; ovary (8-)5–2-locular, superior to

rarely semiinferior; ovules numerous to two

per locule; style usually simple or with

short stylar branches, stigma sometimes

thickened, capitate and/or lobed. Seeds usu-

ally glabrous, rarely winged or arillate;

endosperm reduced to absent; cotyledons

folded. n = 36–46.. . . . .10. bombacaceae.

15 Pollen grains generally spiny, only rarely

smooth. Fruits schizocarpic or loculicidal

or septicidal capsules, rarely baccate or

samaroid. Herbs or shrubs, only rarely

trees; vestitute principally of stellate, lepi-

dote, simple, and unbranched glandular

hairs, sometimes spiny. Leaves simple or

lobed, exceptionally digitate, mostly stipu-

late. Flowers solitary or in cymes, collec-

tively forming spicate to paniculate or even

head-like inﬂ orescences, actinomorphic or

more or less zygomorphic, bisexual or

sometimes unisexual; epicalyx present or

absent, of (2)3 or more bracts, occasionally

persistent; calyx lobes valvate in bud, usu-

ally nectariferous within at base; corolla

tubular to rotate or reﬂ exed; petals ﬁ ve,

convolute in bud; stamens monadelphous;

ﬁ laments few to many, terminating the

staminal column; anthers reniform, 1-celled,

dorsiﬁ xed. Pollen grains often spiny.

Carpels (1-)3 to many, ovary superior; style

exceeding the staminal column, single with

lobed, decurrent or divergent stigmas as

many as the carpels; ovules 1–9 or many in

each carpels, ascending to pendulous. Seeds

reniform, trigonous or turbinate, rarely aril-

late; embryo curved, endosperm present or

absent, cotyledons folded, n = 5–13, 15–17,

19–23, 29, 33, 36–39.. . . . 11. malvaceae.

1. MUNTINGIACEAE

Bayer, M.W. Chase and F. Fay 1998. 3/3. Central and

tropical South America, West Indies.

1.1 MUNTINGIOIDEAE

Leaves serrate. Ovary superior, surrounded by a disc-like

elevation of the receptacle, 7–5-locular. – Muntingia.

1.2 NEOTESSMANNIOIDEAE

Leaves cordate, stipulate. Calyx deeply lobed. Anthers

loculi not conﬂ uent at the apex, opening by longitudi-

nal slits or apical pores. Pollen grains in tetrads. Ovary

inferior, 5-locular (Dicraspidia) or multilocular in

the lower part and 1-locular in the upper part

(Neotessmannia). – Neotessmannia, Dicraspidia.

The basal family in the Malvales.

2. TILIACEAE

A.L. de Jussieu 1789 (including Berryaceae

Doweld 2001, Grewiaceae Doweld et Reveal 2005,

Sparmanniaceae J.G. Agardh 1858). 39/450+. Widely

distributed in tropical regions, especially in tropical

America, Africa, and Southeast Asia. The genus Tilia

(80) extends into the temperate regions of the Northern

Hemisphere. The largest genus Grewia (150) occurs in

tropical Africa, Asia, and Australasia. Corchorus

(about 40 species in Africa and Asia) is one of the few

herbaceous genera in the family.

2.1 TILIOIDEAE

Sepals free. Anthers with the separate locules. Ovary

superior, with 2-many anatropous ovules in each

locule. – apeibeae: Glyphaea, Apeiba, Ancistrocarpus;

enteleae: Entelea, Mortoniodendron, Burretio-

dendron, Sicrea; corchoreae: Corchorus (? including

Oceanopapaver – Tirel et al. 1996; Whitlock et al.

268 Subclass V. DILLENIIDAE

2000), Pseudocorchorus; sparmannieae: Sparmannia,

Clappertonia; lueheeae: Luehea, Trichospermum,

Mollia; tilieae: Tilia, Schoutenia; duboscieae:

Duboscia; desplatsieae: Desplatsia, Vasivaea,

Hydrogaster; grewieae: Grewia, Eleutherostylis,

Tetralix, Lueheopsis; coloneae: Colona, Goethalsia;

triumfetteae: Erinocarpus, Triumfetta, Heliocarpus;

craigieae: Craigia.

2.2 BROWNLOWIOIDEAE

Sepals united into a 3–5th or lobed campanulate

calyx. Anthers with the loculi often conﬂ uent at the apex.

Ovary superior, 1–5-locular; in Brownlowieae carpels

soon free, indumentum often lepidote, n = 10. – bro wn-

lo wieae: Christiana (including Asterophorum and

Tahitia), Brownlowia, Pentaplaris; diplodisceae:

Diplodiscus, Pityranthe (including Hainania),

Jarandersonia; berryeae: Berrya, Carpodiptera,

Pentace.

Tiliaceae are related to the Elaeocarpaceae but differ

markedly in the character of their indumentum, in the

presence of usually well-developed mucilage cavities,

mucilage cells, or canals, ﬁ bers with simple pits, alter-

nate lateral pitting of the vessels, mostly apotracheal

axial parenchyma, usually somewhat more specialized

rays, stratiﬁ ed phloem in young stems, and, as Corner

(1976) notes, also in the anatomy of the seed coat.

3. DIPTEROCARPACEAE

Blume 1825. 13/500. Tropical regions of the Old

World, especially the rain forests of Malesia.

shoreae: Hopea, Shorea, Neobalanocarpus, Upuna;

parashoreae: Parashorea; dryobalaneae:

Dryobalanops; dipterocarpeae: Cotylelobium, Vateria,

Vateriopsis, Vatica, Stemonoporus, Dipterocarpus,

Anisoptera.

Dipterocarpaceae are close to the Tiliaceae and

especially to the Monotaceae. According to Hallier

(1901, 1912), Dipterocarpaceae (but without Monotes)

belong to the Malvales and are related to Elaeocarpaceae

and Tiliaceae.

4. MONOTACEAE

Maury ex Takhtajan 1987. 4/40. Tropical Africa

(Monotes and Marquesia) and tropical South America

(Pakaraimaea and Pseudomonotes).

4.1 MONOTOIDEAE

Flowers with an androgynophore. Sepals imbricate.

Petals longer than the sepals, variously pubescent.

Anthers little or deeply basiversatile. Ovary

(3)4-locular, each locule with two ovules. Trichomes

fasciculate, often glandular. Wood rays uniseriate. –

Monotes, Marquesia.

4.2 PAKARAIMAEOIDEAE

Flowers without androgynophore. Sepals imbricate.

Petals shorter than sepals, glabrous. Anthers deeply

basiversatile. Ovary 4(5)-locular, each locule with

four ovules. Trichomes fasciculate. Wood rays

biseriate. – Pakaraimaea.

4.3 PSEUDOMONOTOIDEAE

Flowers without androgynophore. Sepals valvate.

Petals longer than sepals, glabrous. Anthers imbedded

in thickened connective, with prominent appendage.

Ovary 3-locular, each locule with one subbasal ovule.

Trichomes glandular. – Pseudomonotes.

Monotaceae are closely related to the Tiliaceae

from which they differ mainly in imbricate sepals and

vestured intervessel pores (in Tiliaceae known only for

Schoutenia). Heim (1892) transferred the genus

Monotes to the Tiliaceae, and according to Fries

(1914), Marquesia macroura is related to Schoutenia

in Tiliaceae. According to Bancroft (1933, 1935) the

subfamily Monotoideae serves as a connecting link

with tropical Asiatic members of the Tiliaceae.

Finally, Kostermans (1978) argued for the close

relationships between the genus Pakaraimaea and

Monotes and Marquesia and concluded that they

should be transferred to the Tiliaceae. They are

undoubtedly closer to the Tiliaceae than to the

Dipterocarpaceae. Based on the wood anatomical data,

Gottwald and Parameswaran (1966) suggested raising

the subfamily Monotoideae into a separate family

Monotaceae, and later this was conﬁ rmed by palyno-

logical studies (Maury 1981).

5. SARCOLAENACEAE

Caruel 1881 (including Rhodolaenaceae Bullock

1958, Schizolaenaceae Barnhart 1895). 10/35.

Madagascar.

Sarcolaena, Leptolaena, Xyloolaena, Perriero-

dendron, Eremolaena, Schizolaena, Rhodolaena,

Pentachlaena, Mediusella, Xerochlamys.

Superorder MALVANAE 269

Sarcolaenaceae are related to the Tiliaceae and

especially to Dipterocarpaceae and have many com-

mon features with them in the structure of stem, peti-

ole, and pollen grains (Metcalfe and Chalk 1950;

Dehay 1957; Carlquist 1964; Takhtajan 1997).

Hutchinson (1969) draws attention to a remarkable

feature of most of the genera – the metamorphosis of

the bracteoles, which are united to form an involucre

or epicalyx usually enclosing the fruit – as well as to

another characteristic of the family, the pollinia, con-

sisting of 4 or 16 pollen grains.

6. NEURADACEAE

Link 1831 (including Grielaceae Martynov 1820).

3/10. Arid regions of North Africa, Arabian Peninsula,

Syria, Iraq, northern Iran, Afghanistan, and from

Pakistan to India (Neurada), southern and southeast-

ern Africa (Grielum and Neuradopsis).

Neurada, Grielum, Neuradopsis

Neuradaceae related to the Malvaceae (Hallier

1907, 1912; Airy Shaw 1973; Melikian and Bondar

1996; Bayer 2003).

7. STERCULIACEAE

E.P. Ventenat ex Salisbury 1807 (including

Byttneriaceae R. Brown 1814, Chiranthodendraceae

A. Gray 1887, Dombeyaceae Kunth 1829, Helicteraceae

J.G. Agardh 1858, Hermanniaceae Marquis 1820,

Lasiopetalaceae Reichenbach 1823, Melochiaceae

J.G. Agardh 1858, Pentapetaceae Brechtold et J. Presl

1820, Theobromataceae J.G. Agardh 1858, Triplobaceae

Raﬁ nesque 1838, Triplochitonaceae K. Schumann

1900). 62/1500. Pantropical, extending into subtropical

and warm-temperate regions.

7.1 BYTTNERIOIDEAE

Flowers bisexual or less often polygamous. Petals

mostly present, sometimes very small, scalelike or even

absent (in some Lasiopetaleae and in Fremontodendreae).

Carpels mostly united until dehiscence, rarely free at

maturity. – lasiopet aleae: Hannafordia, Leptony-

chiopsis, Thomasia, Seringia, Keraudrenia, Max wellia,

Lysiosepalum, Lasiopetalum, Guichenotia; herman-

nieae: Hermannia, Melochia, Dicarpi dium, Waltheria;

helmiopsideae: Helmiopsiella, Helmiopsis,

Nesogordonia; byttnerieae: Rulingia, Commersonia,

Byttneria, Ayenia; theobromeae: Glossostemon,

Scaphopetalum, Leptonychia, Abroma, Theobroma,

Herrania, Guazuma; helictereteae: Pterospermum,

Helicteres, Neoregnellia, Kleinhovia, Reevesia,

Ungeria; fremontodendreae: Fre montodendron,

Chiranthodendron; eriolaeneae: Eriolaena;

triplochitoneae: Triplochiton.

7.2 DOMBEYOIDEAE

Flowers usually in axillary cymes or solitary; petals

often persistent, ﬂ at, with apical appendage; pollen

usually sphaeroidal and spinose. – Ruizia, Astiria,

Cheirolaena, Trochetia, Helmiopsiella, Dombeya,

Paradombeya Harmsia, Melhania, Paramelhania,

Pentapetes.

7.3 STERCULIOIDEAE

Flowers mostly unisexual or polygamous, rarely

(in some Tarrietieae) bisexual. Petals absent (present

in Mansonia). Androgynophore present. Carpels

free or at least free at maturity, mostly dehiscent

and not winged (Sterculieae) or indehiscent and

obliquely winged or keeled. – sterculieae: Sterculia,

Brachychiton, Firmiana, Scaphium, Pterocymbium,

Pterygota, Acropogon, Octolobus, Cola; tarrietieae:

Heritiera, Argyrodendron, Tarrietia; mansonieae:

Mansonia, Hildegardia

Sterculiaceae stand close to both the Tiliaceae and

the Bombacaceae and Malvaceae. The boundaries

between these families are however not very clear, and

the taxonomic position of some genera, such as the

paleotropical genus Leptonychia (“The strangely aril-

late seed has a ﬁ brous exotegmen which is neither

Sterculiaceae nor Tiliaceous,” states Corner 1976: 261)

and of the very isolated New Caledonian genus

Maxwellia is still not certain.

Sterculiaceae are very near to the Tiliaceae, which

is conﬁ rmed by the anatomy of vascular system of the

petiole (Dehay 1941, 1942). But the family

Sterculiaceae is very diverse and is characterized by a

great diversity of morphological, anatomical, and

palynological features. In particular, there are so

many differences between the Byttnerioideae and

Sterculioideae that Edlin (1935) proposed to accept

them as two separate families “on account of the struc-

ture of their wood, vegetative organs, ﬂ owers, and

fruit” (p. 6). However, the boundary between these two

groups is not so very clear-cut. For example, on the

basis of the vascular system of the petiole, the tribe

Byttnerieae is closer to the Sterculieae than to the other

270 Subclass V. DILLENIIDAE

tribes of the Byttnerioideae (Gazet du Chatelier 1940a, b).

Besides, as Rao (1952) has shown, the structure of the

androecium is similar for the whole family, and there-

fore there is no reason to separate the Sterculioideae in

a family of their own.

8. DIEGODENDRACEAE

Capuron 1964. 1/1. Madagascar.

Diegodendron.

According to Capuron (1962, 1970), Hutchinson

(1969, 1973), Straka and Albers (1978), and Cronquist

(1980, 1988), the nearest ally of Diegodendraceae is

the family Ochnaceae. This conclusion is based mainly

on the gynobasic style, which is evidently the result of

parallel evolution. The specialized wood anatomy of

Diegodendron supports a close afﬁ nity with the mal-

valean families, particularly with the Sphaerosepalaceae

(Dickison 1988). However, according to Nandi (1998),

Diegodendraceae are closely related to the Bixaceae,

and Thorne (2006) places them between the Bixaceae

and Cochlospermaceae.

9. SPHAEROSEPALACEAE

van Tieghem ex Bullock 1959 (Rhopalocarpaceae

Hemsley 1903). 2/17. Madagascar.

Dialyceras, Rhopalocarpus.

Very closely related to the Diegodendraceae. Both

of these families exhibit deﬁ nite afﬁ nities to the Sarco-

laenaceae.

10. BOMBACACEAE

Kunth 1822. 29/c.300. Pantropical, especially in rain

forests of South America, above all in Brazil.

durioneae: Neesia, Durio, Camptostemon,

Coeloste gia, Boschia, Kostermansia, Cullenia,

Scleronema, Cavanillesia; matisieae: Quararibea,

Ochroma, Matisia, Patinoa, Bernoullia, Septotheca,

Phragmo theca, Huberodendron; catostemateae:

Aguiaria, Catostemma; bombaceae: Bombax,

Eriotheca, Adan sonia, Pseudobombax, Pachira

(including Rhodog naphalon, Rhodognaphalopsis, and

Bombacopsis) Gyranthera; ceibeae: Ceiba,

Spirotheca, Chorisia, Neobuchia.

Very close to the Malvaceae.

11. MALVACEAE

A.L. de Jussieu 1789 (including Hibiscaceae J.G. Agardh

1858, Philippodendraceae Endlicher 1841, Plagiantha-

ceae J.G. Agardh 1858). 111/1500–1600. Essentially

cosmopolitan, but best developed in the tropics.

kydieae: Julostylis, Dicellostyles, Nayariophyton,

Kydia; hibisceae: Decaschistia, Radyera, Hibiscus,

Papuodendron, Talipariti, Kosteletzkya, Fioria,

Wercklea, Abelmoschus, Hibiscadelphus, Senra,

Symphyochlamys, Megistostegium, Perrierophytum,

Humbertiella, Macrostelia, Helicteropsis, Humber-

tianthus, Cenocentrum, Lagunaria, Urena, Malachra,

Peltaea, Phragmocarpidium, Rojasimalva (includ-

ing Lopimia), Pavonia, Malvaviscus, Anotea;

gossypieae: Cienfuegosia, Cephalohibiscus, Lebron necia,

Hampea, Thespesia, Gossypioides, Kokia, Gossypium,

Alyogyne; malveae: Anoda, Periptera, Horsfordia,

Bakeridesia, Bastardiastrum, Wissadula, Tetrasida,

Pseudabutilon, Bastardia, Bastardiopsis, Abutilon,

Herissantia, Neobaclea, Meximalva, Corynabutilon,

Briquetia, Hochreutinera, Dirhamphis, Gaya,

Billieturnera, Sidastrum, Sida, Krapovickasia,

Rhynchosida, Robinsonella, Dendrosida, Fryxellia,

Allowissadula, Allosidastrum, Neobrittonia, Batesi-

malva, Malvella, Lecanophora, Cristaria, Astero-

trichion, Plagianthus, Gynatrix, Hoheria, Lawrencia,

Sidasodes, Sidalcea, Eremalche, Iliamna, Malvastrum,

Modiolastrum, Modiola, Calyculogygas, Callirhoe,

Calyptraemalva, Napaea, Monteiroa, Lavatera, Malva,

Althaea, Navaea, Malope, Malacothamnus, Phymosia,

Alcea, Kitaibelia, Anisodontea, Kearnemalvastrum,

Acaulimalva, Urocarpidium, Fuertesimalva, Palaua,

Nototriche, Tarasa, Sphaeralcea, Andeimalva.

Malvaceae are so closely related to the Bomba-

caceae, that from time to time the proposal arises to

merge these two families. Edlin (1935) transferred

the tribe Hibisceae together with the genus Kydia to

the Bombacaceae, but as Hutchinson (1967: 538)

noted, “Malvaceae without the great genus Hibiscus

would be like a horse without a tail.” The most

archaic tribe is Malopeae, in which the carpels are in

two or more superposed and spirally arranged cycles.

All other tribes have carpels in a single cycle.

The tribe Ureneae, probably the most advanced, has

twice as many style branches as carpels, which may

be due to a splitting of the styles, just as the anthers

are split and unilocular in the whole family

(Hutchinson 1967: 538).

Superorder MALVANAE 271

12. BIXACEAE

Kunth 1822. 1/5. Tropical America and West Indies.

Bixa.

13. COCHLOSPERMACEAE

Planchon 1847. 2/15–20. Tropical and subtropical

regions of America, western and central tropical Africa,

southern and southeastern Asia and northern Australia.

Cochlospermum, Amoreuxia.

Very close to the Bixaceae.

14. CISTACEAE

A.L. de Jussieu 1789 (including Helianthemaceae G.

Meyer 1836). 8/180–200. Mostly temperate and sub-

tropical regions of the Northern Hemisphere, espe-

cially the Mediterranean and eastern North Africa,

with some in West Indies and South America.

cisteae: Cistus, Halimium, Crocanthemum,

Tuberaria, Helianthemum (including Atlanthemum?),

Fumana; hudsonieae: Hudsonia; lechidieae: Lechea.

15. TEPUIANTHACEAE

Maguire et Steyermark 1981. 1/7. Venezuela, Brazil, and

Colombia; lowland savannas and Roraima sandstone.

Tepuianthus.

Very close to the Thymelaeaceae.

16. THYMELAEACEAE

A.L. de Jussieu 1789 (including Aquilariaceae

R. Brown ex A.P. de Candolle 1825, Daphnaceae

Ventenat 1799, Gonystylaceae van Tieghem 1896,

Phaleriaceae Meissner 1841). 48/800. Subcosmopolitan

but concentrated mainly in tropical Africa and

Australia, with local centers in the Mediterranean

region and western, eastern, and Southeast Asia.

Classiﬁ cation after Domke (1934), revised by B.E.

Herber (2003).

16.1 OCTOLEPIDOIDEAE (GONYSTYLOIDEAE)

Trees, rarely shrubs. Inﬂ orescences usually determinate;

bracts minute or absent. Floral tube not articulated,

persistent; sepals (3)4–5(6), valvate or imbricate; petals

4–40, free or sometimes slightly fused at base, rarely

forming a ﬂ eshy annulus, inserted at receptacle, or absent.

Stamens inserted at receptacle; ﬁ laments slender,

sometimes broader at base; anthers straight, reﬂ exed,

peltate or horseshoe-shaped. Pollen grains 8–60-porate,

or rarely (Octolepis) 3–6-porate, and reticulate. tectate-

columellate. Nectary disc wanting. Gynoecium of

(2)3–5, rarely 6–8 carpels, with elongate, ﬁ liform,

wiry, contorted style, occasionally accompanied by a few

small processes (parastyles) around the base; stigma

small, capitate to punctiform. Fruits 1–9-seeded,

usually loculicidally dehiscent or (Aetoxylon) 1-seeded,

indehiscent. Seeds large, without chalazal fold, usually

with thin dorsal aril arising from the ﬂ eshy funicle or

(Amyxa and Aetoxylon) not arillate; endosperm wanting.

– octolepideae: Octolepis; microsemmateae: Lethedon

(Microsemma); solmsieae: Deltaria, Solmsia,

Arnhemia; gonystylideae: Gonystylus, Amyxa,

Aetoxylon.

16.2 THYMELAEOIDEAE

Trees or shrubs (sometimes lianas), rarely perennial

herbs, very rarely annuals; interxylar phloem present

in some genera. Inﬂ orescences usually indeterminate,

occasionally with involuctum. Sepals 4–5(6), connate

into a campanulate or cylindric tube, often broader

around the ovary, imbricate (valvate in Lagetta); petals

well-developed, entire or divided, ringlike, scalelike,

reduced to small, ﬂ eshy glands, obscure or lacking.

Stamens equal or double the number of calyx lobes

(at most 10) or very rarely (Pimelea) reduced to two

or even one. Pollen grains 8–60-porate and with

more or less clearly expressed crotonoid ornamenta-

tion. Nectary disc annular, cupular, composed of

scales, or lacking. Gynoecium of two carpels. Ovary

2-locular (Peddiea, Synandrodaphne, Oreodendron,

and Phaleria) or more often 1-locular. Fruits indehis-

cent, nutlike, baccate, or drupaceous, or more rarely

capsule. Seeds sometimes appendiculate, with or

without endosperm. – synandrodaphneae:

Synandrodaphne (Gilgiodaphne); aquilarieae:

Aquilaria, Gyrinops; phalerieae: Peddiea, Phaleria;

dicranolepideae: Lachnaea, Lagetta, Goodallia,

Funifera, Lophostoma, Gnidia, Enkleia, Linostoma,

Stephanodaphne; daphneae: Jedda, Dicranolepis,

Synaptolepis, Craterosiphon, Linodendron, Daphnopsis,

Schoenobiblus, Ovidia, Dirca, Daphne, Wikstroemia,

Rhamnoneuron, Edgeworthia, Thymelaea, Diarthron,