Takhtajan Armen. Flowering Plants

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Superorder RHAMNANAE 433

Rhamnaceae, and Thorne (2000, 2006) includes

Dirachmaceae in the Rhamnales. According to Baas,

Jensen and Smets (2001), Dirachma has many com-

mon anatomical features with Rhamnaceae. However

Dirachmaceae markedly differ from the Rhamnaceae

and related families in the presence of epicalyx, the

lack of a nectary disc, and structure of gynoecium and

fruit. The caducous glands of Dirachmaceae are not

homologous with the persistent receptacular disc-nec-

taries of Rhamnaceae and Elaeagnaceae (Smets 1986;

Smets and Cresens 1988).

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434 Subclass VI. ROSIDAE

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A. Takhtajan, Flowering Plants, 435

Trees, shrubs, subshrubs, perennial or annual herbs.

Vessels with scalariform or simple perforations. Sieve-

element plastids usually of S-type. Nodes trilacunar,

multilacunar or unilacunar. Leaves simple or com-

pound, alternate, opposite or verticillate. Stomata of

various types. Flowers in various kinds of inﬂ orescences

or sometimes solitary, mostly bisexual or less often uni-

sexual, actinomorphic or more or less zygomorphic,

often characterized by the secondary pollen presenta-

tion. Corolla often sympetalous. Stamens usually ﬁ ve,

often attached to the corolla tube. Anthers tetrasporan-

giate, opening longitudinally. Pollen grains 2-celled or

3-celled, colpate, colporate or porate. Tapetum secre-

tory or (some Dipsacales) amoeboid. Microsporogenesis

simultaneous. Pollen grains 2- or 3-celled, 3-colporate or

of derived types. Nectary disc mostly present. Gynoecium

of 2–5 (up to 12) united carpels; stylodia free or more or

less connate; ovary mostly inferior. Ovules anatropous,

unitegmic, mostly tenuinucellate. Endosperm cellular

or more often nuclear. Fruits of various types. Seeds with

small or large embryo; endosperm copious to scanty,

rarely absent.

Probably originated from a Cunoniales-like saxifra-

ganaean ancestor.

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436 Subclass VII. ASTERIDAE

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Superoder CORNANAE

Order 106. DESFONTAINIALES

Trees or shrubs, less often subshrubs or herbs. Vessels

mostly with scalariform perforations (sometimes

with numerous bars), lateral pitting scalariform to

alternate. Fibers mostly with bordered pits, often

septate. Rays heterogeneous. Axial parenchyma

apotracheal to scanty paratracheal, often wanting.

Sieve-element plastids of S-type. Nodes trilacunar or

less often unilacunar or multilacunar. Leaves alternate,

opposite or less often verticillate, simple, entire or less

often lobed or dentate, stipulate or more often estipu-

late. Stomata paracytic or more often anomocytic.

Flowers in inﬂ orescences of various types or rarely

solitary, bisexual or sometimes unisexual, actinomor-

phic or sometimes more or less zygomorphic. Sepals

(3)4–5(-12), valvate or imbricate, free or more often more

or less connate. Petals (3)4–5(-12), valvate, imbricate

or contorted, free or connate into a short tube. Stamens

(3)4–5 or 8–10 or more, sometimes numerous (even up to

200 in Carpenteria); ﬁ laments free, basally connate or

adnate to the corolla tube; anthers opening longitudinally.

Tapetum secretory. Microsporogenesis simultaneous.

Pollen grains 2-celled, mostly 3-colporate, sometimes

2–3-colpate. Intrastaminal nectary disc present or less

often wanting. Gynoecium of (2)3–5 or less often up to

12 united carpels; stylodia free or more often connate

into a style with the stigmatic branches or lobed or capi-

tate stigma; ovary superior or more often semi- inferior

or inferior, with 1-several or many ovules per locule.

Ovules mostly anatropous, usually unitegmic, tenuinu-

cellate, or rarely crassinucellate, usually with endothe-

lium. Female gametophyte of Polygonum- or Allium-type.

Endosperm cellular, with terminal haustoria. Fruits

septicidal or loculicidal capsules or less often baccate

or drupaceous. Seeds exotestal, rarely (Vahliaceae)

endotestal, with straight embryo surrounded by ﬂ eshy

endosperm or less often without endosperm.

The Escalloniaceae, and related families are usually

included in Rosales sensu amplissimo or in Cunoniales

or even in the family Saxifragaceae s. 1. However,

this group of families differs from Saxifragaceae,

Cunoniaceae, and related families and is much closer to

the Cornales (see Huber 1963; Dahlgren 1975, 1980,

1983, 1989 [who even included them in the Cornales];

Takhtajan 1987; Soltis et al. 1990; Thorne 1992a, b,

2006; Xiang et al. 1993). From the saxifraganaean and

rosanaean families are differ not only morphologically

(unitegmic and tenuinucellate ovules) but also in the

presence of iridoid compounds (Jensen et al. 1975;

El-Naggar and Beal 1980). Its distinctness is also sup-

ported by serological studies (Grund and Jensen 1981).

Key to Families

1 Ovules bitegmic.

2 Vessel elements long (sometimes very long),

with exclusively scalariform perforations that

have 16–71 (occasionally up to 111) thin bars;

lateral pitting extremely rare and conﬁ ned to

overlapping end walls (when present typically

occur in uniseriate ﬁ les). Fibers with bordered

pits. Rays of homocellular, uniseriate rays of

upright cells and heterocellular, bi- and multise-

riate rays. Axial parenchyma sparse and predom-

inantly apotracheal diffuse and paratracheal

scanty. Evergreen tree up to 15 m tall. Nodes tri-

lacunar with three traces. Leaves alternate, oppo-

site or verticillate, coarsely and bluntly

glandular-serrate, estipulate. Stomata anomo-

cytic. Marginal veins are not ﬁ mbriate. Unicellular

T-shaped trichomes occur on both the pedicels

Superoder CORNANAE 437

and ﬂ owers. Flowers in terminal, subumbellate

panicles. Sepals spirally arranged, and the three

lower ones, which are larger than the other two,

completely enclose the other organs in the bud.

Calyx-tube adnate to base of ovary, lobes ﬁ ve,

imbricate, deciduous. Petals ﬁ ve, inserted below

5-lobed disc, clawed, imbricate. Connectives are

not prolonged. Stamens ﬁ ve, alternate with disc

lobes, ﬁ laments slender. Pollen grains 2-celled,

(3)4–5-colporate. Ovary superior to semi-inferior,

conical, 5-lobed, 5-locular, with two ovules per

locule; style spirally twisted, 5-grooved. Ovules

crassinucellate, with nucellar cap, funicular obtu-

rator and ruminate arillode and without endothe-

lium. Fruits broadly ovoid, coriaceous, 5-locular,

loculicidal capsules. Seeds large, glossy, with a

broad “aril”, hilar scar elongated, exotestal; coty-

ledons large. Contains proanthocyanidins (from

cortex), urolic acid; n = 25. . . . . . 4. ixerbaceae.

2 Vessels with simple perforations. Leaves oppo-

site, simple, entire, ovate to linear, estipulate.

Stomata anomocytic. Annual herbs rather densely

pubescent by multicellular glandulose hairs.

Nodes unilacunar. Flowers small, paired, in axil-

lary cymose inﬂ orescences, bisexual, actinomor-

phic. Sepals ﬁ ve, connate into hemispherical tube

adnate to the ovary; lobes valvate. Petals ﬁ ve,

free, shorter than the calyx lobes, obovate-spatulate,

imbricate. Stamens ﬁ ve, alternipetalous, inserted

on the margin of the epigynous nectary disc; ﬁ la-

ments free, subulate; anthers dorsiﬁ xed, introrse.

Pollen grains 2-celled, 3-colporate. Gynoecium

of two or three united carpels, with two or three

thick, divergent stylodia terminated by capitellate

stigma; ovary inferior, 1-locular, with numerous,

anatropous, bitegmic, tenuinucellate ovules

attached to the large placentas hanging from the

apex of the locule. Fruits more or less globose,

loculicidal capsules, 2–3-valved at the top, with

persistent calyx. Seeds numerous, minute, oblong,

appendaged; exotestal cells more or less elon-

gated; endotesta a layer of cells with well-devel-

oped U-thickening; embryo large, straight;

endosperm scarce, of two cell layers. Contain iri-

doids n = 6, 9. . . . . . . . . . . . . . .10. vahliaceae.

1 Ovules unitegmic.

3 Trees, shrubs, subshrubs or perennial herbs.

4 Corolla not sympetalous. Stem and petiole

without endodermis.

5 Flowers usually bisexual, rarely polyg-

amo-dioecious.

6 Nectary disc usually present.

7 Pericycle without a ring of lactifer-

ous sacs and without a composite and

continuous ring of sclerenchyma.

Trees and shrubs. Leaves alternate or

rarely (Polyosma) opposite or subop-

posite, estipulate. Hairs unicellular

and thick-walled, glandular-shaggy

with uniseriate stalks of variable

length and aspheroid or (Quintinia)

peltate glands. Vessels with scalari-

form perforations that have many

bars (up to 125 in some species of

Polyosma); lateral pitting transitional,

opposite or alternate. Fibers with bor-

dered pits. Rays heterogeneous. Axial

parenchyma apotracheal, diffuse and/

or diffuse-in-aggregates. Nodes (at

least in Escallonia) unilacunar.

Leaves entire, serrate or dentate,

often glandular-serrate. Flowers in

terminal or axillary cymes, small to

rather large (Anopterus), actinomor-

phic. Sepals 4–6, free or connate into

a short tube. Petals 4–6, free, imbri-

cate or valvate. Stamens mostly ﬁ ve,

free, alternate with the lobes of

nectary disc. Pollen grains 3-colpo-

rate or (Quintinia) 5-colporate.

Gynoecium of 2–5 carpels; stylodia

connate into a short or elongate style

with capitate, 2–5-lobed stigma;

ovary superior to inferior, usually

with numerous ovules on axile or

less often (Polyosma, Anopterus)

parietal placentas. Fruits septicidal

or loculicidal capsules or (Polyosma,

Abrophyllum) berries. Seeds small

or minute; embryo small, surrounded

by copious ﬂ eshy endosperm.

Contain decarboxylated iridoids,

ﬂ avonols, n = 12 (Escallonia).

. . . . . . . . . . . . 1. escalloniaceae.

7 Pericycle with a ring of laticiferous

sacs consisting of vertically elon-

gate cells arranged in longitudinal

rows and ﬁ lled with white, friable

438 Subclass VII. ASTERIDAE

contents and with a composite and

continuous ring of sclerenchyma.

Trichomes glandular with short

sunken stalks and unicellular heads.

Vessels with scalariform perfora-

tions. Leaves alternate, large, long-

acuminate, subserrate. Stomata with

a pair of small cells nearly circular

in outline. Flowers in terminal or

axillary cymes, yellowish. Calyx

tube very short, lobes 6–5, small,

decieduous. Petals 6–5, valvate,

spreading, deciduous. Stamens 6–5,

inserted on the margin of the incon-

spicuous nectary disk; ﬁ laments

very short. Gynoecium of ﬁ ve car-

pels; stigma sessile, 5-lobed; ovary

superior, 5-locular, with numerous

axile ovules. Fruits a small, black,

many-seeded berry, corsned by the

stigma. Seeds small, with deeply

latticed testa; embryo very small.

. . . . . . . . . . .3. abrophyllaceae.

6 Nectary disc absent. Prostrate, gla-

brous shrublets with robust, ﬂ exuous

stems emitting short ascending,

densely leafy branches. Vessels with

scalariform perforations that have

12–14 bars. Leaves alternate, sessile,

semiamplexicaul, simple, entire but

apically minutely tridentate, more or

less glaucous above, obscurely veined,

estipulate. Flowers small, solitary at

apex of branchlets; subsessile, ebrac-

teate, actinomorphic, 5-merous as to

the perianth and androecium. Sepals

small, shortly connate bellow, imbri-

cate, persistent. Petals relatively large,

slightly clawed, contorted, ﬂ eshy or

not. Stamens alternipetalous; ﬁ laments

subulate; anthers small, ovoid, sub-

didymous, extrorse. Pollen grains

3-colporate, rugulate-reticulate, with

fractioned, partial tectum with inter-

rupted muri on the intercolpium.

Gynoecium of three carpels; style

short, with 3-lobed, capitate stigma;

ovary superior, 3-locular, with many

(30–50) anatropous ovules per locule.

Fruits small, many-seeded, loculicidal

and valvular capsules, borne on a

short, erect pedicel; the valves ﬁ nally

separating from the axis, to which the

seeds long remain attached after the

capsule has dehisced. Seeds black,

shining, with ﬂ eshy and oily

endosperm. . . . . . . . .5. tribelaceae.

5 Flowers unisexual, dioecious. Vessels

usually with simple perforations, even in

the primary xylem; lateral pitting alter-

nate, pits more or less vestured. Fibers

with simple pits. Rays mostly heteroge-

neous. Nodes unilacunar.

8 Petals valvate. Small trees or shrubs.

Vessels with simple perforation or

rarely with simple and scalariform

perforations, with 1–2 bars. Fibers not

septate. Rays essentially heteroge-

neous. Axial parenchyma absent.

Leaves alternate, entire, pinnately

veined, coriaceous, estipulate. Stomata

anomocytic. Flowers small, in small

axillary panicles. Male ﬂ owers: calyx

4-angled, 4-lobed. Petals four, oblong-

ovate, free; stamens four; ﬁ laments

very short, anthers basiﬁ xed, oblong-

linear, acuminate. Pollen grains 3-col-

porate, reticulate. Nectary disc

pulvinate, obtusely 4-angled; rudimen-

tary gynoecium subulate. Female ﬂ ow-

ers: calyx tube obtusely 4-angled, teeth

four, remote; petals four, staminodia

absent; nectary disc hemispherical.

Gynoecium of two carpels; stylodia

minute, recurved, with decurrent stig-

mas; ovary semi-inferior, 2-locular,

with one ascending, campylotropous

ovule in each locule. Fruits small, com-

pressed, didymous drupes with two

compressed, 1-seeded, crustaceous,

thin-walled pyrenes. Seeds linear-

oblong, with membranous testa and

ﬂ eshy, orange or brownish endosperm;

embryo large, with inferior radicle,

n = 16. . . . . . . . . . .6. kaliphoraceae.

8 Petals imbricate. Small trees or shrubs,

sometimes (Grevea bosseri) lianas.

Vessels with simple perforation. Fibers

Superoder CORNANAE 439

with bordered pits, septate (Grevea).

Rays heterogeneous (Montinia) or

homogeneous (Grevea). Axial paren-

chyma scanty paratracheal. Leaves

alternate, subopposite or opposite,

entire, coriaceous (Montinia) or mem-

branous (Grevea), estipulate. Stomata

anomocytic (Montinia) or anisocytic

(Grevea). Flowers in terminal or axil-

lary cymes or panicles, or solitary and

terminal (female), small, actinomor-

phic, unisexual, 4–5-merous. Male

ﬂ owers: sepals 3–5, connate into cup-

ular or ﬂ attened, entire or shortly 3–5-

lobed tube; petals 3–5, alternipetalous,

thick; stamens 3–5, alternipetalous;

ﬁ laments short, thick; anthers rather

large, ellipsoid, dorsiﬁ xed, extrorse.

Pollen grains 3-colporate reticulate;

rudimentary gynoecium absent or

minute; nectary disc ﬂ at, discoid.

Female ﬂ owers: sepals 4–5, adnate to

ovary, limb shortly tubular, entire or

minutely 4–5-toothed; petals 4–5,

ﬂ eshy, deciduous; staminodia 4–5;

nectary disc epigynous, ﬂ eshy,

4–5-angled. Gynoecium of two car-

pels; style short, thick, persistent, stig-

mas 2-ﬁ d or shortly 2-lobed; ovary

inferior to semi-inferior, 2-locular,

with numerous (Montinia) or 4–6

ovules per locule; ovules pendulous,

anatropous. Fruits 2-locular and locu-

licidal, dehiscent (Montinia) or inde-

hiscent (Grevea) capsules with few

seeds and tipped by persistent style.

Seeds compressed and winged all

around and auriculate at the point of

attachment (Montinia) or subglobose,

and not winged; exotesta ligniﬁ ed,

periclinal walls thickened; endosperm

more or less developed in Grevea

(Krach 1976), or copious in Montinia

(Nemiro vich-Danchenko 2000).

n = 34 (Montinia). 7. montiniaceae.

4 Corolla sympetalous.

9 Ovules tenuinucellate. Evergreen shrubs

or small trees; plants usually bitter tast-

ing. Vessels with scalariform perfora-

tions; lateral pitting is generally absent

owing to the solitary nature of vessels

and tends toward alternate when present.

Fibers are tracheids. Rays entirely unise-

riate and comprise axially elongated or

upright cells only. Axial parenchyma

vasicentric, scanty with some diffuse.

Leaves opposite, rather small, simple,

asymmetrical, entire or dentate, with

glands on lower surface or stipules pres-

ent. Stomata anomocytic. Flowers in

few-ﬂ owered terminal cymes or solitary

and terminal, two bracteolate, bisexual,

slightly zygomorphic, (4)5(-8)-merous

as to the perianth. Sepals connate, lobes

valvate or weakly imbricate, persistent.

Petals yellow, connate into a very short

tube, lobes spreading, imbricate. Stamens

two, inserted near the base of corolla,

alternate with the adaxial and lateral

lobes; ﬁ laments short, stout; anthers large,

extrorse, with broad connective and two

undulately plicate, latrorse, twisted loculi

that open longitudinally. Pollen grains

3-colporate. Gynoecium of two carpels;

style short, thick, stigma cup-like, with

two broad lobes; ovary almost quite infe-

rior, imperfectly 2-locular, with two

intruded parietal placentas that are subcon-

tiguous in the center; ovules numerous,

ascending, anatropous, with endothelium.

Fruits septicidal and valvular capsules,

with persistent calyx. Seeds numerous,

minute, laterally compressed, smooth, with

minute straight embryo and copious, ﬂ eshy

endosperm.. . . . . . . .8. columelliaceae.

9 Ovules crassinucellate. Erect or sprawl-

ing evergreen shrubs or rarely small trees

with somewhat angular, opposite

branches, glabrous or with a few simple

hairs. Pith parenchymatous. Vessels with

scalariform perforations; perforation

plates very oblique, the bars ﬁ ne and

numerous from 15 to 65; lateral pitting

partly scalariform, partly small, round,

bordered. Fibers with bordered pits. Rays

uniseriate, composed of upright cells.

Axial parenchyma mainly diffuse, but

also some isolated strands associated

440 Subclass VII. ASTERIDAE

with the vessels (Mennega 1980). Sieve-

element plastids of S-type. Leaves oppo-

site, simple, petiolate, spinulose-dentate

like those of Ilex aquifolium, pinnately

veined, coriaceous, connected at the

base by stipular line. Stomata anomo-

cytic. Flowers solitary in the axils of the

leaves near the apices of the branchlets

or in few-ﬂ owered cymes, large, bisex-

ual, actinomorphic or nearly actinomor-

phic, 5-merous, with leafy or sepal-like

bracts. Sepals basally connate, strongly

imbricate, equal or subequal, persistent.

Corolla much longer than the calyx,

nearly cylindrical, ﬂ eshy, lobes imbri-

cate or contorted, scarcely spreading.

Stamens ﬁ ve, inserted just below the

corolla throat, equal; ﬁ laments much

shorter than the anthers; anthers bas-

iﬁ xed, narrowly triangular to nearly

oblong, deeply cordate at the base,

introrse. Pollen grains 3-colporate, with

a thick aperture membrane, rugulate tec-

tum surface with small perforations, and

elongate pores (Hoc and Bravo 1984).

Gynoecium of ﬁ ve united carpels; style

long, slender, with capitate, scarcely

dilated, obscurely 5-lobed stigma, only

slightly expanded, persistent; ovary

superior, cylindrical, 5-locular at the

base and 1-locular at the apex, with ﬁ ve

parietal placentas with many ovules;

placentas mushroom-shaped on section,

united inside in the basal half of the

ovary. Ovules anatropous, with

endothelium. Fruits berries with a per-

sistent calyx. Seeds many, obliquely

ellipsoid or ovoid, slightly angular,

outer walls of testa pectic, other walls

ligniﬁ ed; embryo straight, minute;

endosperm copious, starchy. Producing

iridoid glycosides loganin and loganic

acid (Houghton and Ming 1985). n = 7.

. . . . . . . . . . . . . 9. desfontainiaceae.

3 Annual herbs. Leaves alternate, simple,

somewhat ﬂ eshy, the radical leaves spatu-

late-oblanceolate, entire to dissected, when

dissected, basically palmatiﬁ d (the upper

members stagshornlike), estipulate. Small,

densely glandular-hairy annual herbs, much

branched from the base, thinly pilose all

over. Vessels with simple perforations.

Flowers very small, in terminal, cymose,

paniculate inﬂ orescences, bisexual, actino-

morphic, 5-merous. Sepals narrow, deeply

lobed, basally connate into a very short

tube, adnate to the lowest third of the ovary;

lobes valvate. Petals free, narrowly elliptic,

spreading. Stamens ﬁ ve, alternate with the

petals, free; ﬁ laments subulate; anthers

small, rounded-ellipsoidal, dorsiﬁ xed, ver-

satile, introrse to latrorse. Pollen grains

3-colporate, reticulate. Epigynous disc

absent. Gynoecium of two united carpels,

with free slender divergent stylodia termi-

nated by capitate stigma; ovary 2-locular,

three-quarters superior, shortly adnate to

the calyx tube at the base, with one sub-

basal axile ascending, anatropous ovule

per locule. Fruits semi-inferior, sub-

didymous, loculicidal capsules. Seeds

small, erect, with minute, straight embryo

and copious endosperm; seed coat 3–4

layered.. . . . . . . . . . . .2. eremosynaceae.

1. ESCALLONIACEAE

R. Brown ex Dumortier 1829 (including Anopteraceae

Doweld 2001; Polyosmaceae Blume 1851). 6/150. The

largest genus Polyosma (60) is distributed from eastern

Himalayas through Southeast Asia to New Guinea,

tropical Australia, and New Caledonia, Escallonia

(about 40) in South America (especially Andes),

Quintinia (25) from Malesia to Australia, New Zealand,

and New Caledonia, Cuttsia (1), and Anopterus (2) are

endemic to Australia, Forgesia (1) to Reunion and

Valdivia (1) to Chile.

Anoptereae: Anopterus; FORGESIEAE:

Forgesia, Quintinia; escallonieae: Escallonia,

Valdivia; polyosmeae:

Polyosma.

The family, even in its narrow sense, as deﬁ ned, is

rather heterogeneous.

2. EREMOSYNACEAE

Dandy 1959. 1/1. Southwestern Australia.

Eremosyne

Furthermore, Al-Shammary and Gornall (1994)

suggested a close relationship between Eremosyne and

Superoder CORNANAE 441

Escallonia based on trichome anatomy. According to

Soltis and Soltis (1997), Eremosynaceae are the sister

to the Escalloniaceae.

3. ABROPHYLLACEAE

Nakai 1943. 2/2. Eastern Australia.

Cuttsia, Abrophyllum

Related to the Escalloniaceae, but differing in the

structure of pericycle and Abrophyllum and Cuttsia

both have clusters of small, unligniﬁ ed cells in the

mesophyll that look like little white raphide bundles

(Hils 1985).

4. IXERBACEAE

Grisebach 1854. 1/1. Northern New Zealand (lowland

and lower montane forests from lat. 35°30′ to a little

south of 3 8°).

Ixerba

Vegetative anatomy of Ixerba is very primitive.

5. TRIBELACEAE

Airy Shaw 1965. 1/1. Temperate South America (from

southern Chile to Tierra del Fuego).

Tribeles

Related to the Escalloniaceae, differing from them

in the absence of nectary disc, in contorted petals,

extrorse anthers, persistent seed-bearing column of the

fruits, and the obscurely veined, glaucescent leaves

(“very distinctive in appearance, and unlike any others

known to the writer,” states Airy Shaw [1965: 269]), as

well as in seed coat anatomy (Nemirovich-Danchenko

and Lobova 1998).

6. KALIPHORACEAE

Takhtajan 1996. 1/1. Madagascar.

Kaliphora

Capuron was the ﬁ rst who noticed the uncornaceous

features of Kaliphora, which were later conﬁ rmed

by Eyde (1988). Kaliphora is related to both the Montin-

iaceae. There are many palynological similarities

between Montinia, Grevea, and Kaliphora (Hideux and

Ferguson 1976; Ferguson 1977; Ferguson and Hideux

1978). The wood anatomy of Kaliphora resembles that

of Montinia and Grevea (Bakolimalala-Ramamonjiarisoa

1989). However, Kaliphora differ from the Montiniaceae

s. str. in many respects, including free stylodia, solitary

pendulous ovule, and drupaceous fruits. According to

Eyde (1988: 310), Kaliphora «has commissural stig-

mas: receptive tissue extends downward in the septal

radii not in the radii of the dorsal carpel bundles as is the

case in the comparable Cornaceae (that is, in the

Nyssoideae: only they have well-deﬁ ned style branches».

Also, as Eyde points out «Insertion of the ovule is lower

– about midlevel in the cavity – in Kaliphora than in true

cornads and the cushionlike epigynous nectary of true

cornads is absent. Furthermore, the seeds develop with

their micropyle turned down».

7. MONTINIACEAE

Nakai 1943. 2/4 Southern Africa (Montinia) and tropi-

cal West and East Africa and Madagascar (Grevea).

Montinia, Grevea

Related to the Escalloniaceae, and in Hutchinson’s

system (1967, 1969, 1973) they are included in them.

However, as early as 1891 Kerner von Marilaun and later

Nakai (1943) and independently Takhtajan (1954) and

Milne-Redhead (1955) considered both genera a separate

family, which was later accepted by Capuron (1969),

Airy Shaw (in Willis 1973), Krach (1976), Dahlgren

(1980, 1983, 1989), Carlquist (1989) and Thorne (1992a,

b). Capuron (1969) also proposed to include in this family

Kaliphora, which was accepted by Dahlgren (1980, 1983)

and later by Carlquist (1989) and Thorne (1992), who

also added Melanophylla. Palynologically Montinia,

Grevea, and Kaliphora are very similar (Hideux and

Ferguson 1976; Ferguson and Hideux 1978). Montiniaceae

are considerably more advanced than Escalloniaceae.

According to Carlquist (1989), wood of Montinia resem-

bles that of Kaliphora and Grevea, but Grevea is more

distant from Montinia than is Kaliphora.

8. COLUMELLIACEAE

D. Don 1828. 1/4. Andes of South America from

Colombia to Bolivia.

Columellia

Probably related to the Escalloniaceae and differing

from them mainly in absence of the nectary disc,

weakly sympetalous corolla, slightly zygomorphic

442 Subclass VII. ASTERIDAE

ﬂ owers, androecium of two stamens with broad con-

nective and contorted anthers, and thick integument.

A great variety of families have been proposed as rela-

tives of Columellia (Columelliaceae). They include

Oleaceae, Gesneriaceae, Ericales, Ebenales, Genti-

anales, Scrophulariales, Myrtales, Saxifragales, and

even Cucurbitaceae. The most recent treatments favor

inclusion in Cornales (Dahlgren 1975, 1983, 1989),

Resales s. 1. (Cronquist 1981, 1988; Thorne 1983),

Saxifragales s. 1. (Hallier 1908, 1911, 1912; Takhtajan

1980), or Hydrangeales (Takhtajan 1987; Thorne

1992a, b). Hallier (1908, 1911, 1912) included

Columellia in Saxifragaceae-Philadelpheae. According

to Stern et al. (1969: 70), it is clearly impossible to

assemble an array of data that would afﬁ rm unequivo-

cally the relationships of Columelliaceae with any one

of the several families to which it has been allied. In

their opinion «Perhaps its nearest relatives are in the

Escalloniaceae» (p. 70).

9. DESFONTAINIACEAE

Endlicher 1841. 1/5. The Andes from Costa Rica to

Cape Horn.

Desfontainia

Closely related to the Columelliaceae (Hallier 1908,

1911, 1912). Thorne (1992a, b, 2000, 2006) placed

the Desfontainiaceae in his Cornales close to the

Columelliaceae. A relationship between Desfontainia

and Columellia is also supported by the wood anatomy

(Mennega 1980). Backlund and Donoghue (1996) even

included Desfontainia in Columelliaceae. However,

there are also many differences in ﬂ oral morphology.

10. VAHLIACEAE

Dandy 1959. 1/8. Tropical and South Africa, Madagascar,

and from upper Egypt and Iran to northwestern India.

Vahlia (Bistella)

Usually included in the Saxifragales, but Vahliaceae

differ from them in their placentas are pendulous,

ovules are tenuinucellate, and micropyle is formed

by the inner integument. Besides, mature seeds do

not show any traces of the raphe (Krach 1976, 1977).

And what is even more important, Vahlia contains

iridoids (Al-Shammary 1991). According to rbsL and

18S rDNA sequence data, Vahlia is well separated

from Saxifragales and included in the same clade

which contains Montinia (Morgan and Soltis 1993;

Soltis and Soltis 1997). According to Al-Shammary

and Gornall (1994) Vahlia resembles Montinia also in

thin-walled, eglandular, uniseriate trichomes with a

smooth cuticle, opposite, estipulate leaves. Probably

belong to the Hydrangeales. However, Vahlia differs

from the typical members of Desfontainiales in

bitegmic ovules. Systematic position of Vahlia is still

uncertain.

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