Takhtajan Armen. Flowering Plants

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Superorder CELASTRANAE 413

Secretory cells with tanniniferous contents present in

the unligniﬁ ed tissues of leaves and stem. Leaves all

basal, ﬂ at, long petiolate, the blades palmately veined,

ovate, reniform, or orbicular, entire, with cuneate,

truncate, or cordate bases; leaf venation campylodro-

mous. Stomata on the abaxial leaf surface. Flowers

protandrous, solitary, on a long peduncle, terminal on

scapiform stems with one or (in section Fimbripetalum)

many sessile and reduced leaves, often with a clasping

base. Sepals persistent. Petals showy, larger than

sepals, sessile, or clawed, imbricate, persistent.

Stamens dehiscing over the gynoecium in sequence

before moving outwards; anthers ventriﬁ xed, versatile,

extrorse, the connective is broad with a short protru-

sion. Staminodia ﬁ ve, candelabra-like, consist of three

or many ﬁ liform divisions with glandular-appearing

apices. Nectar secretion from the bases of the stamin-

odes. Pollen grains 2-celled, 3-colporate, 4-colporate

and syncolpate, with thin exine and ﬁ ne columellae,

reticulate. Seeds minute (in Parnassia palustris about

100–2,000 in capsules – Hultgärd 1987), oblong,

thinly, with a winglike testa; endotegmic cells with

U-shaped thickening; endosperm scanty or absent,

embryo straight, cylindrical, n = 7, 8, 9. – Parnassia.

8.2 LEPUROPETALOIDEAE

Ovules unitegmic. Diminutive annual, more or less

succulent herbs, usually forming small hemispherical

tufts. Leaves both basal and cauline, alternate or

subopposite, spatulate, entire, the blade constricted

into a broad petiole; leaves brown-streaked from tan-

nins; leaf venation acrodromous. Stomata on both

leaf surface. Flowers small and inconspicuous, either

appearing sessile in the basal rosette or terminal at the

ends of short, leafy, and angled stems. Petals white,

smaller than sepals, sometimes absent or of (4)5

linear, bractlike, submicroscopic structures inserted

on the rim of the ﬂ oral cup. Stamens very short;

anthers bisporangiate at anthesis (Simmons 2004),

Staminodia external to the stamens, dilated distally.

Seeds cylindrical, blackish, with a short beak on

one end, the surface reticulate; endotestal cell walls

much thickened, endosperm scanty, n = 23. –

Lepuropetalon.

9. HUACEAE

A. Chevalier 1947. 2/3. Tropical Africa.

Hua, Afrostyrax.

The afﬁ nities of Huaceae have long been obscure

(Soltis et al. 2005). According to Zhang and Simmons

(2006), “Huaceae are best supported as the sister group

of Oxalidales, suggesting that it should be placed

within that order.”. Here I follow to recent version of

Thorne system (2006) in inclusion of Huaceae in the

Celastrales.

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Superorder SANTALANAE

Order 102. SANTALALES

Trees, shrubs, woody lianas, or perennial herbs,

mostly hemiparasitic and attaching to the roots or

stems of other plants. Vessels with simple or some-

times scalariform perforations; lateral pitting scalari-

form to alternate. Fibers with bordered or simple

pits. Rays heterogeneous or homogeneous. Axial

parenchyma mostly apotracheal. Sieve-element plas-

tids of Ss- or (Misodendraceae) So-type. Nodes trila-

cunar or unilacunar, rarely pentalacunar. Leaves

alternate or opposite, simple, entire, mostly estipu-

late. Stomata of various types. Inﬂ orescences axillary

or terminal. Flowers mostly rather small, bisexual or

unisexual, usually actinomorphic, often apetalous or

in the most reduced groups even naked. Calyx often

very reduced or absent. Petals free or basally connate,

mostly valvate, seldom absent. Stamens usually

isomerous and opposite the petals; ﬁ laments free or

sometimes adnate to the petals (very rarely to the

sepals) or connate into a column; anthers basiﬁ xed,

tetrasporangiate, disporangiate, or monosporangiate,

opening longitudinally or by apical pore or trans-

verse slit. Tapetum secretory. Microsporogenesis

simultaneous. Pollen grains 2-celled or sometimes

3-celled, with various types of apertures (from

3- colpate to pantoporate). Gynoecium of 2–3(-4)

united carpels, with short or rarely long style; ovary

superior to inferior, mostly 1-locular or partitioned

only at the base, with the ovules pendulous from a

columnar placenta that extends upward beyond the

basal partitions. Ovules anatropous, epitropous,

hemitropous, or orthotropous, bitegmic to unitegmic

and usually tenuinucellate, but more often ategmic,

sometimes not differentiated from the placenta and

mostly characterized by clearly expressed reduction

of nucellus. Female gametophyte of Polygonum- or

sometimes of Allium-type, rarely (Thesium) of

Adoxa-type. Endosperm cellular or less often

helobial. Fruits of various types. Seeds mostly

without seed coat, usually with well-developed

endosperm; embryo terete and lug-like. Contain

triglycerides with C18 acteylenic acids, triterpenic

sapogenins.

Probably related to Celastrales (Hutchinson 1973;

Dahlgren 1983).

Key to Families

1 Ovules well differentiated from placenta (except in

Exocarpos of Santalaceae). Ovary superior or infe-

rior. Endosperm cellular or helobial.

2 Plants usually terrestrial. Calyx present, at least

in female ﬂ owers, but usually greatly reduced in

Schoepﬁ a, and minute or almost absent in

Opiliaceae.

3 Leaves alternate. Ovary superior or inferior.

Stamens opposite the petals.

4 Nectary disc present. Petals usually

present.

5 Stamens not united into a column.

Gynoecium of 2–5 carpels.

6 Nectary disc annular, more or less

adherent around the base of ovary.

Trees, woody lianas, and shrubs, rarely

subshrubs, sometimes root parasites

(Schoepﬁ a and Ximenia), sometimes

with schizogenous resin glands and/or

branched laticifers. Nodes mostly trila-

cunar with three traces. Vessels usually

with simple perforations, but in some

genera perforations scalariform with

6–10 bars (up to 20 or more in

Heisteria); lateral pitting scalariform to

alternate. Fibers with bordered or more

often simple pores. Rays heterogeneous

with long or short ends. Axial paren-

Superorder SANTALANAE 417

chyma apotracheal to paratracheal.

Flowers usually small, 3–6-merous, in

axillary panicles, racemes, heads, or

(Erythropalum) very slender compound

dichasia, sometimes solitary in the

axils, usually bisexual or rarely unisex-

ual (dioecious). Calyx small, more or

less cupular, mostly 3–6-toothed, often

accrescent in fruit. Petals 3–6, usually

valvate or rarely imbricate, free or

basally connate, seldom forming a long

tube. Stamens as many as and opposite

the petals or more often 2–3 times as

many, but apparently always in one

cycle; ﬁ laments free or sometimes

adnate to the corolla, very rarely adnate

to calyx; anthers 4-locular, opening

longitudinally or sometimes by termi-

nal valves. Pollen grains 3-colpate,

3-colporate or 3–8-porate. Gynoecium

of (2)3(-5) carpels. Stigma 2–5-lobed;

ovary free or more or less embedded in

nectary disc, 2–5-locular at the base,

but mostly 1-locular above, with 1

ovule in each locule or semilocule, pen-

dulous from the top of the free central

placenta; several genera, e.g.,

Anacolosa, have a complete placental

column attached to the top of ovary;

ovules mostly anatropous, bitegmic,

unitegmic, or ategmic, tenuinucellate

or very rarely (Ximenia americana)

crassinucellate. Endosperm cellular (as

in Strombosia) or helobial (as in Olax).

Fruits usually 1-seeded drupes or nuts,

often subtended or enclosed by accres-

cent and colorful calyx, the pericarp

ﬂ eshy, thin, the endocarp woody. Seeds

with green small or minute embryo

near tip of copious, oily (and sometimes

also starchy) endosperm; cotyledons

2–6; seed coat thin, crushed. n = 12, 13,

16, 19, 20. . . . . . . . . . . .1. olacaceae.

6 Nectary disc of separate glands or parts

only united at the base but not adherent

to ovary, alternating with stamens. Trees

and shrubs, sometimes lianoid, usually

or always hemiparasitic. Vessels with

simple perforations; lateral pitting alter-

nate. Fibers with bordered or rarely sim-

ple pits. Rays homogeneous. Axial

parenchyma apotracheal, usually dif-

fuse. Cystoliths present. Flowers small,

in axillary or cauliﬂ orous inﬂ orescences

(spikes, racemes, umbels, or panicles),

bisexual or (Gjellerupia and Agonandra)

dioecious, 4–5-merous. Calyx small and

inconspicuous, cupular or with 4–5

small lobes or teeth. Petals free or con-

nate below or less often for more than

half of their length, valvate, in female

ﬂ owers usually wanting. Stamens as

many as and opposite petals; ﬁ laments

free or borne on petals or on corolla

tube; anthers 4-locular, opening longi-

tudinally; rudimentary gynoecium

present. Pollen grains 3-colporate,

microechinate. Stigma small or capi-

tately expanded or stigma sessile; ovary

free or half-sunken in nectary disc, supe-

rior, 1-locular, with a single (very rarely

2) ovule pendulous from the summit of

the columnar free- central placenta or

(Agonandreae) the ovule basal and erect.

Ovules pendulous, anatropous (or

orthotropous ? – Corner 1976), uniteg-

mic and tenuinucellate or not divided into

nucellus and integument (Agonandra).

Endosperm cellular. Fruits drupaceous;

the pericarp thin, usually yellow to

orange, the mesocarp ﬂ eshy, the endo-

carp woody or crustaceous. Seeds large,

with rather small embryo and copious,

oily and starchy endosperm; seed coat

thin-walled, crushed; cotyledons (2)

3–4, n = 10. . . . . . . . . . . 2. opiliaceae.

5 Stamens united into a column around the

style with as many sessile anthers in a

whorl at the top. Trees. Vessels with sim-

ple perforations; lateral pitting alternate.

Fibers with simple or (Ongokea) dis-

tinctly bordered pits. Rays heteroge-

neous. Axial parenchyma typically

apotracheal. Flowers in short, axillary

inﬂ orescences racemes (Harmandia) or

panicles, usually bisexual or dioecious.

Calyx 4–5-dentate or -lobed, small at

anthesis but very much enlarged and

418 Subclass VI. ROSIDAE

persistent in fruit. Petals 4–5 in bisexual

ﬂ owers, or 6–8 in female ones, free, val-

vate. Nectary disc extrastaminal, annular

(Harmandia) or lobed between petals.

Stamens 4 or (Ongokea) 5; anthers 2-loc-

ular, opening from apex to base by a

membranous ﬁ nally reﬂ exed valve. Pollen

grains (3)4(5)-porate. Gynoecium of 3–2

carpels, with elongated style between the

staminal column or (Harmandia) stigmas

sessile; ovary 2- or 1-locular, with 1–3

ovules pendulous from the top of the free-

central placenta. Ovules not divided into

nucellus and integument. Fruits drupes,

more or less enclosed by the enlarged

calyx; pericarp coriaceous, endocarp

woody. Seeds with oily endosperm and

very small embryo near its tip. Ongokea

has polyacetylenes. . .3. aptandraceae.

4 Nectary disc absent or present in female

ﬂ owers.

7 Ovary superior. Trees with scattered secre-

tory canals containing a yellow liquid.

Vessel elements very long with oblique,

scalariform perforation plates that have

11–43 (mostly 19–24) bars; lateral pitting

scalariform. Rays high, uniseriate-hetero-

geneous. Axial parenchyma scanty para-

tracheal and diffuse. Leaves alternate,

apparently simple, remotely and minutely

crenulate, pinnately veined, transverse

veins numerous and parallel; the lower

surface provided with hairs of a unique,

thick-based type; petiole with a pulvinus at

the top, suggesting that the leaves may be

unifoliolate rather than simple; stipules

small, deciduous. Stomata anomocytic.

Flowers small, in axillary, pendulous cat-

kinlike racemes, bisexual, actinomorphic.

Sepals ﬁ ve, free or nearly so, open in bud,

persistent. Petals ﬁ ve, free, small, imbri-

cate. Fertile stamens ﬁ ve, oppositipetalous,

short, erect in bud, free from each other but

ﬁ laments slightly adnate to the base of pet-

als; anthers tetrasporangiate, the outer

(abaxial) loculi opening by an outwardly

recurved valve, the inner (adaxial) ones

opening by an inwardly recurved valve;

staminodia ﬁ ve, oppositisepalous, elon-

gate, long-exerted, linear, very densely and

shortly pubescent above the glabrous base,

in-ﬂ exed at the apex and with an abortive

anther at the tip. Pollen grains 3- colporate.

Nectary disc wanting. Gynoecium of 3

(-4) united carpels; stylodia free, widely

separate, short, with a very small, terminal

stigma; ovary 1- locular, with a slender

free-central placental column and 6(-8)

anatropous, epitropous ovules pendulous

from the top of the column. Fruits

3(4)-valved coriaceous 1-seeded capsule,

silky-ﬁ brous within, subtended by persis-

tent reﬂ exed sepals. Seeds large, pulvini-

form, often with six radiating ribs above,

with small, straight embryo near the

edge of the copious, slightly ruminate

endosperm. . . . . . . 5. medusandraceae.

7 Ovary inferior. Trees or shrubs with stellate

hairs (as in Couleae, Olacaceae). Vessels

with scalariform (Octoknema) or simple

perforations (Okoubaka); lateral pitting

opposite to alternate. Fibers with simple

pores, rarely septate. Rays heterogeneous

with long ends (Octoknema) or homoge-

neous (Okoubaka). Axial parenchyma apo-

tracheal, diffuse or (Octoknema) absent.

Stomata anomocytic. Flowers usually in

axillary racemes, spines, or panicles, uni-

sexual, 5-merous. Sepals valvate, mostly

very reduced, in male ﬂ owers obsolete.

Petals more or less connate, valvate, persis-

tent, the outer surface of the petals densely

covered by hairs. Stamens free, oppositipe-

talous; ﬁ laments short and broad; anthers

short, introrse. Pollen grains 3-colporate.

Rudimentary gynoecium in male ﬂ owers

with 5-lobed disc at the base. Staminodia in

female ﬂ owers without anthers or

(Okoubaka) with sterile anthers. Nectary

disc in female ﬂ owers lobed. Gynoecium of

three carpels; style very short and thick,

with three broad, reﬂ exed, irregularly lobu-

late lobes; ovary inferior, in lower part

3-locular, more or less 1-locular above with

three ovules at the apex of a threadlike,

free-central placenta that reaches and adnate

to the top of the ovary. Ovules anatropous,

unitegmic. Fruits drupaceous, 1-seeded.

Superorder SANTALANAE 419

Seeds with small embryo and copious,

slightly ruminate endosperm; radicle much

longer than cotyledons; cotyledons up to 6.

n = 36.. . . . . . . . . . . . 4. octoknemaceae.

3 Leaves opposite or less commonly alternate,

sometimes reduced to scales, rarely (Ionidium)

margins with spines. Stomata mostly paracytic.

Hemiparasitic terrestrial small trees, shrubs,

and perennial herbs usually on roots of other

plants or rarely (as Dendrotrophe) on branches,

sometimes thorny or xeromorphic. Cuticular

epithelium common (cuticle waxes as rodlets);

epidermal cells scleriﬁ ed, with druses. Vessels

with simple perforations; lateral pitting usually

alternate. Fibers with bordered or simple pores.

Rays heterogeneous or homogeneous. Axial

parenchyma usually rather scanty, apotrachel,

or paratracheal. Nodes unilacunar. Flowers

small, often greenish, in various kinds of inﬂ o-

rescences (often small dichasium in axil of each

bract), rarely solitary in leaf axil (as in Thesium),

bisexual or unisexual (monoecious or dioe-

cious). Calyx extremely reduced, inconspicu-

ous. Petals (3)4–5(-8), free or forming a

valvately lobed, often ﬂ eshy cup or tube.

Stamens as many as and opposite the petals;

ﬁ laments often adnate to base of petals; anthers

4-locular or rarely (Exocarpos) 2- locular, open-

ing longitudinally of by single apical pore.

Pollen grains 3-aperturate, with apertures of

various types. Lobed nectary disc mostly pres-

ent (absent in Thesieae), surrounding or seated

on the ovary or lining the lower part of ﬂ oral

tube. Gynoecium of (2)3(-5) carpels, with ter-

minal style and lobed or capitate stigma; ovary

superior to inferior, completely 1-locular or

with basal partitions, with straight or spirally

coiled free-central placenta bearing 1–4, less

often 4–5 pendulous ovules. Ovules anatropous

or less often hemitropous, or orthotropous,

unitegmic and tenuinucellate, ategmic (Mida

and Santalum) or (Exocarpos) more or less

undifferentiated into nucellus and integument.

Endosperm cellular or (Mida and Santalum)

helobial. Fruits drupaceous, nuts, or baccate.

Seeds without a differentiated testa, with

straight embryo surrounded by copious, ﬂ eshy,

oily, or (Thesium) starchy endosperm; seed coat

absent. n = 5–7. . . . . . . . . . . .6. santalaceae.

2 Hemiparasitic shrublets almost always on

Nothofagus spp., with radially expanded hausto-

rial region and stout twigs; shoot apex aborts

annually and one or more lateral branches con-

tinue vegetative growth in a sympodial fashion

in the next growing season. Vessel elements very

short, with simple perforations; lateral pitting

alternate to scalariform. Fibers very short and

broad, with bordered pits. Rays uniseriate, homo-

geneous. Axial parenchyma vasicentric. Leaves

alternate, small, sometimes scalelike. Stomata

mostly paracytic. Flowers very small, in axillary

catkinlike compound racemes or spikes, sub-

tended by a bract, dioecious, but very occasion-

ally bisexual (Skottsberg 1913); male ﬂ owers

without perianth of two or three stamens around

small, lobed nectary disc; anthers at least ini-

tially 2-locular, monothecous, opening by apical

tangential slit. Pollen grains 4–12- colpate.

Female ﬂ owers apetalous, but with three sepals

basally adnate to the ovary and free at the top,

they fused with the ovary along their midregion

and extended sideways in a winglike fashion,

having three deep grooves that rub the length of

the ovary; from each groove protrudes a short

bristlelike staminodium that is lengthening into

long plumose bristle during maturation of the

fruit. Gynoecium of three carpels; style very

short, stout, with 3-lobed stigma; ovary essen-

tially superior (Kuijt 1969), 1-locular, with

three ovules pendulous from the top of short,

free, central placental column, each hanging

down into a basal pocket of ovarian cavity;

ovules anatropous, reduced, not differentiated

into nucellus and integument. Female gameto-

phyte Polygonum-type. Endosperm cellular,

with elongate chalazal haustorium. Fruits small,

1-seeded nuts crowned the strongly accrescent

staminodia covered by numerous long,

unicellular trichomes and serving as an adapta-

tion for wind dispersal and anchoring while the

seeds germinate. Seed without testa or testa rudi-

mentary (M. punctulatum), with straight embryo

surrounded by oily green endosperm; seed coat

absent. . . . . . . . . . . . . . . . .7. misodendraceae.

1 Ovules embedded in placenta, placentalike body, or

in the base of the ovary and scarcely differentiated

from them. Ovary inferior, usually 1-locular.

Endosperm cellular. Seeds without testa.

420 Subclass VI. ROSIDAE

8 Flowers usually bisexual, only rarely unisexual.

Hemiparasitic brittle shrublets on tree branches

(or on epiphytes), less often terrestrial shrubs,

lianas, or even (Nuytsia) trees on host roots, with

a single or several haustoria at ends of epicortical

roots, occasionally almost Cuscuta-like in habit.

Stem often dichasially branched, but without

nodal construction. Vessels with simple perfora-

tions; lateral pitting alternate. Fibers short to very

short, with simple or clearly bordered pits. Rays

slightly to strongly heterogeneous. Axial paren-

chyma apotracheal. Nodes unilacunar. Leaves

mostly opposite, less frequently alternate or ver-

ticillate, always simple, sometimes reduced to

scales. Stomata mostly paracytic. Flowers rather

large and brightly colored to small, in various

types of inﬂ orescences the basic unit of which is

a triad or simple dichasium, rarely (Ixocactus)

solitary, bisexual or rarely unisexual (plant dioe-

cious), actinomorphic or (in African members)

with a strong tendency to zygomorphic, fre-

quently red or yellow, entomophilous or ornitho-

philous. Calyx represented by toothed or lobed

rim around the summit of ovary. Petals (3-)5–6(-

9), valvate, free or often connate below into a

tube, sometimes nectariferous basally within.

Stamens as many as and opposite to petals; ﬁ la-

ments often adnate to petals; anthers 4-locular or

sometimes 2-locular or 1- locular opening longi-

tudinally. Pollen grains 3- or seldom 4-(5)-aper-

turate, rarely (Atkinsonia) inaperturate, trilobate

or triangular. Nectary disc present or wanting.

Gynoecium of 3–4-carpels, with very short to

much elongated style and small stigma; ovary

1- locular, with commonly 4–12 ovules embed-

ded in the erect, free-central placenta or the basal

tissue of the ovary, rarely (Lysiana) ovary 4-locu-

lar with an axile placentas; each ovule consists

essentially of monosporic, 8-nucleate female

gametophyte of Polygonum-type. Suspensor

highly elongate. Fruits usually laticiferous ber-

ries or drupes, mostly 1-seeded, less often

2–3-seeded; rarely fruits dry and indehiscent.

Seeds more or less covered with viscous mate-

rial, embryo green, rather large, at least some-

times without obvious radicle; endosperm

copious, starchy, rarely wanting at maturity.

Contain triterpenes (loranthin, betulin, etc.),

rather large amounts of tannins, alkaloids,

saponins, and fatty acids, n = 8–12 (Wilson et

Calvin 2006).. . . . . . . . . . . . .8. loranthaceae.

8 Flowers unisexual, generally small and incon-

spicuous. Calyx obsolete. Ovules two. Female

gametophyte bisporic.

9 Leaves opposite. Hemiparasitic, brittle shrub-

lets on trees without epicortical roots; haustoria

( sometimes almost like mycelium) penetrating

and ramifying in host tissues; aerial branding

often pseudodichotomous and often with nodal

constructions. Vessels with simple perforations;

lateral pitting alternate. Fibers very short, with

simple or evidently bordered pits. Rays more or

less heterogeneous. Axial parenchyma apotra-

cheal. Leaves opposite, often reduced to scales.

Stomata mostly paracytic. Flowers very small,

not brightly colored (usually yellow or green),

more or less sessile, mostly in spikelike often

branching inﬂ orescences with (1)3-ﬂ owered

dichasium in axil of each bract, unisexual

(monoecious or dioecious), anemophilous or

entomophilous. Petals small, valvate, usually

four in male ﬂ owers and three in female ﬂ ow-

ers, often reduced to merely teeth or bumps on

ovary rim. Stamens opposite and often adnate

to petals; tetrasporangiate and opening longitu-

dinally or reduced to one or two microsporan-

gia and opening by apical pores or slits or

transversely partitioned and opening by a num-

ber of transverse slits, or the anthers conﬂ uent

into synandrium (Viscum album, Korthalsella

opuntia). Pollen grains usually spheroidal, tria-

perturate or in some African Viscum species

(3-)4–6-aperturate, in one African Viscum spe-

cies (5-)7–10-rugorate. Gynoecium of 3–4 car-

pels; style very short with a small stigma; ovary

inferior, usually 1-locular, with massive pla-

centa (mamelon) bearing two ategmic ovules

usually completely reduced to only a female

gametophyte. Female gametophyte bisporic, of

Allium-type. Suspensor very short or absent.

Cleavage of zygote transverse. Fruits

1(2)-seeded berries, sometimes explosive.

Seeds with viscid tissue at one end; embryo

rather large, green, embedded in starchy green

endosperm. Chemistry is similarity of that of

Loranthaceae, except for toxic polypeptides

(visco-toxins and lectins), which present only

in Viscaceae, n = 10–15(-17). . . 9. viscaceae.

Superorder SANTALANAE 421

9 Leaves alternate or sometimes opposite

(Lepidoceras and juvenile plants of Eubrachion).

Small hemiparasitic shrubs on woody plants,

sometimes (spp. of Antidaphne) with epicortical

roots. Vessels with simple perforations. Leaves

foliaceous or (in mature plants of Eubrachion)

peltate and scalelike. Flowers small, mostly

sessile, ebracteolate, in catkinlike spikes or

racemes, often with deciduous scale leaves;

sometimes female ﬂ owers solitary in leaf axils,

unisexual (dioecious or monoecious), 2–4-mer-

ous, without calyx. Aborted organs of opposite

sex wanting. Petals small, two or four (or rarely

none) in male ﬂ owers, two or three in female

ﬂ owers. Stamens free, opposite petals; ﬁ la-

ments often very short; anthers tetrasporangiate,

2-locular, opening longitudinally. Pollen grains

3-colporate, echinate or rarely (Eubrachion)

shallowly reticulate and lacking any spines.

Gynoecium of three or ﬁ ve carpels with a short,

stout style surrounded by a capitate or crestlike

stigma, sometimes shallowly lobed; ovary infe-

rior or semi- inferior, 1- locular and contains a

small, central ovarian papilla (mamelon) bear-

ing two extremely reduced ovules without

clearly deﬁ ned nucellus and integument and

almost completely reduced to only a female

gametophyte. Female gametophyte bisporic of

Allium-type. Fruits 1-seeded berries. Seeds

extremely slippery on account of large amount

of associated viscin concentrated at its radicular

pole; embryo rather large, with two cotyledons

or (Lepidoceras peruvianum) undifferentiated

and without cotyledons; endosperm green or

white (absent in Lepidoceras), n = 10, 13.

. . . . . . . . . . . . . . . . . . .10. eremolepidaceae.

1. OLACACEAE

A. L. de Jussieu ex R. Brown 1818 (including

Cathedraceae van Tieghem 1900, Coulaceae van

Tieghem 1900, Erythropalaceae Planchon ex Miquel

1856, Harmandiaceae van Tieghem 1900, Heisteriaceae

van Tieghem 1900, Schoepﬁ aceae van Tieghem 1900,

Strombosiaceae van Tieghem 1900, Tetrastylidiaceae

van Tieghem 1900, Ximeniaceae Horaninow 1834).

25/200–250. Widespread in tropical and subtropical

regions.

1.1 ANACOLOSOIDEAE

Ovules with one or two integuments, rarely (Ximenia

americana) without integuments, anatropous, with

dorsal raphe; micropyle facing upward. Calyx often

enlarged with ripening of fruit. Ovary superior. –

couleae: Coula, Maburea, Minquartia, Ochano stachys;

heisterieae: Heisteria, Chaunochiton; ximenieae:

Ximenia; anacoloseae: Scorodocarpus, Engomegoma,

Phanerodiscus, Brachynema, Diogoa, Tetrastylidium,

Cathedra, Anacolosa, Strombosia, Strombosiopsis.

1.2 ERYTHROPALOIDEAE

Ovules with one integument, anatropous, with dorsal

raphe. Calyx not enlarged. Ovary embedded in nectary

disc. – Erythropalum.

1.3 OLACOIDEAE

Ovules without integument, anatropous, with dorsal

raphe; micropyle facing upwards. Calyx more or less

enlarged. – Curupira, Ptychopetalum, Douradoa,

Olax, Dulacia, Malania.

1.4 SCHOEPFIOIDEAE

Ovules with one integument, with dorsal raphe. Calyx

inconspicuous, not enlarged. Ovary embedded in

nectary disc. Corolla sympetalous. – Schoepﬁ a.

2. OPILIACEAE

Valeton 1886 (including Cansjeraceae J.G.Agardh

1858). 10/36–72. Widespread in tropical and subtropi-

cal regions, especially in Asia; Agonandra (10) the

only American genus.

opilieae: Lepionurus, Cansjera, Melientha,

Champereia, Opilia, Urobotrya, Rhopalopilia,

Pentarhopalopilia, Gjellerupia; agonandreae:

Agonandra.

Very close to the Olacaceae. According to Baas

et al. (1982), Opiliaceae might be considered a direct

derivative from Olacaceae through Ximenia.

3. APTANDRACEAE

Miers 1853. 3/6. Southeast Asia (Harmandia), tropical

Africa (Ongokea and Aptandra zenkeri), and tropical

South America (three other species of Aptandra).

Aptandra, Ongokea, Harmandia.

Very close to Olacaceae.

422 Subclass VI. ROSIDAE

4. OCTOKNEMACEAE

Solereder 1908. 2/6. Tropical Africa.

Octoknema, Okoubaka.

Very close to Olacaceae.

5. MEDUSANDRACEAE

Brenan 1952. 1/2. Equatorial West Africa.

Medusandra.

Close to Olacaceae and especially to

Octoknemaceae.

6. SANTALACEAE

R. Brown 1810 (including Anthobolaceae Dumortier

1829, Arjonaceae van Tieghem 1898, Canopodaceae

K. Presl 1851, Osyridaceae Raﬁ nesque 1820,

Thesiaceae Vest 1818). 37/450. Widely distributed, but

mostly tropical and subtropical regions; best developed

in relatively dry areas.

santaleae: Choretrum, Spirogardnera, Leptomeria,

Phacellaria, Dufrenoya, Scleropyrum, Jodina,

Cervantesia, Buckleya, Omphacomeria, Pyrularia,

Osyris, Geocaulon, Comandra, Cladomyza, Nanodea,

Myoschilos, Acanthosyris, Santalum, Rhoiacarpos,

Amphorogyne, Daenikera, Colpoon, Dendromyza,

Elaphanthera, Mida, Dendrotrophe, Kunkeliella,

Nestronia; thesieae: Osyridocarpos, Thesidium,

Thesium, Austroamericium, Arjona, Quinchamalium;

anthoboleae: Anthobolus, Exocarpos.

Close to the Olacaceae and originated from them

(Hallier 1912; Fagerlind 1948; Agarwal 1963; Kuijt

1968, 1969).

7. MISODENDRACEAE

J. Agardh 1858. 1/11. Cool-temperate South America

from the Strait of Magellan north to about 33° south-

ern latitude.

Misodendrum.

Close to the Santalaceae, especially to South

American genera Arjona and Quinchomalium

(Skottsberg 1935), but differ in the absence of the peri-

anth of the male ﬂ owers, and the absence of petals in

female ﬂ owers, and male ﬂ owers with monothecous

anthers and a rudimentary testa in the seed, and

probably originated from them (Hallier 1912). Kuijt

(1968, 1969) derives Misodendraceae from Olacaceae.

According to Carlquist (1985), wood anatomy of

Misodendraceae is similar to that of Psittacanthus and

other Loranthaceae, disregarding the unique features

found in Misodendrum.

8. LORANTHACEAE

A. L. de Jussieu 1808 (including Dendrophthoaceae

van Tieghem 1898, Elytranthaceae van Tieghem 1898,

Gaiadendraceae van Tieghem 1898, Nuytsiaceae van

Tieghem 1898, Psittacanthaceae Nakai 1952).

75/c.1000. Mostly in tropical and subtropical regions,

especially in Southern Hemisphere.

nuytsieae: Nuytsia; lorantheae: Gaiadendron,

Panamanthus, Atkinsonia, Desmaria, Amylotheca,

Decaisnina, Lampas, Lepeostegeres, Thaumasianthes,

Cyne, Loxanthera, Peraxilla, Alepis, Lysiana, Trilepidea,

Macrosolen, Elytranthe, Lepidaria, Loranthus,

Helicanthera, Moquiniella, Tristerix, Dactyliophora,

Amyema, Psittacanthus, Amyema, Ileostylus,

Dendropthoe, Baratranthus, Scurulla, Emelianthe,

Pedistylis, Plicosepalus, Oncella, Kingella, Trithecanthera,

Trithecanthera, Taxillus, Vanwykia, Pedistylis, Oliverella,

Oncocalyx, Phragmanthera, Septulina, Tapinanthus,

Agelanthus, Globimetula, Struthanthus, Benthamina,

Erianthemum, Tolypanthus, Papuanthes, Diplatia,

Distrianthes, Tetradyas, Bakerella, Helicanthes,

Sogerianthe, Englerina, Tupeia, Notanthera, Tripod-

anthus, Muellerina, Psittacanthus, Oryctanthus,

Cladocolea, Oryctina, Phthirusa, Dendropemon,

Aetanthus, Ligaria, Actinanthella, Berhautia, Cecarria,

Ixocactus, Socratina, Spragueanella.

Probably derived directly from the Olacaceae

(Hallier 1912), possibly from Chaunochiton-like

ancestor (Kuijt 1968, 1969). According to Kuijt,

Loranthaceae, contrary to tradition, cannot have

evolved from Santalaceae (“The ﬂ ower of

Loranthaceae is dichlamydous and can scarcely

have a monochlamydous ancestor in Santalaceae.”

[1969: 80]).

9. VISCACEAE

Batsch 1802 (including Arceuthobiaceae van Tieghem

1897, Bifariaceae Nakai 1952, Ginalloaceae van