Takhtajan Armen. Flowering Plants

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Superorder SANTALANAE 423

Tieghem 1899, Phoradendraceae H. Karsten 1860).

7/450. Very widely distributed, but mostly in tropical

and subtropical regions; all genera except Arceuthobium

occur either in the Old or the New World; Arceuthobium

is restricted to the Northern Hemisphere.

phoradendreae: Phoradendron, Dendrophthora,

Korthalsella, Ginalloa; arceuthobieae: Arceuthobium;

visceae: Notothixos, Viscum.

Closely allied to the Loranthaceae. According to

Kuijt (1968, 1969), Viscaceae may be traceable to

Santalaceae plants similar to present-day epiphytic

parasites such as Phacellaria.

10. EREMOLEPIDACEAE

Nakai 1952 (including Lepidocerataceae Nakai

1952). 3/12. From Isia Chiloe (Chile) north along the

Andes and the lower, eastern areas of South America

to Venezuela and Colombia; only Antidaphne vis-

coldea reaches through Central America as far as

Chiapas (Mexico); Eubrachion and Antidaphne

wrightii are represented on both Hispaniola and

Puerto Rico; the former also occurs on Jamaica, the

latter in eastern Cuba.

Antidaphne (including Eremolepis), Eubrachion,

Lepidoceras.

Kuijt (1988) recognizes the distinct possibility of

an early separation of Eremolepidaceae from a prelo-

ranthaceous stock. The afﬁ nities to Loranthaceae have

also been stated by various authors, including Bhandari

and Vohra (1983), who mention the striking similari-

ties between the epicortical roots and secondary haus-

toria of the more primitive eremolepidaceous genus

Antidaphne and the loranthacous genus Notanthera.

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Superorder BALANOPHORANAE

Order 103. CYNOMORIALES

Terrestrial reddish-brown parasitic herbs with

branched, thick, ﬂ eshy, underground rhizome. Stem

simple, ﬂ eshy, bearing numerous, alternate, deltoid,

scalelike leaves. Haustorial contacts are made by

means of numerous small lateral roots on the rhizome

and even with the base of the ﬂ owering stem, which

fuses with the host root. Food reserve is starch. Vascular

system reduced. Stomata vestigial, anomocytic.

Flowers minute, numerous, on short peduncles, inter-

spersed with the scalelike leaves in terminal clavate or

cylindrical inﬂ orescence with ﬂ eshy axis, polygamous,

apetalous, entomophilous. Calyx of (1)4–5 (rarely

6–8) strap-shaped, linear-spatulate sepals, free or con-

nate at the base. Stamen single, at the base adnate to

the perianth; anthers dorsiﬁ xed, versatile, tetrasporan-

giate, introrse, opening longitudinally. Tapetum secre-

tory. Microsporogenesis simultaneous. Pollen grains

2-celled, 3-colporate, reticulate. Male ﬂ owers with

426 Subclass VI. ROSIDAE

1–2 bright yellow nectaries-pistillodia. Gynoecium

monocarpellate (Terekhin and Nikiticheva 1976), with

a thick and long stylodium, which forms a long canal

lined with conducting tissue and has a conduplicate

upper part; ovary inferior, unilocular. with 1 pendu-

lous, submarginal ovule. Ovule pendulous, submar-

ginal, orthohemitropous, unitegmic with thick

integument, almost crassinucellate. Female gameto-

phyte of Polygonum-type. Endosperm cellular. Fruits

1-seeded berries (Doweld 2000). Seeds with copious

endosperm and minute, lentiform, undifferentiated

embryo; seed coat of four or ﬁ ve layered; cells of

exotesta covered with thick layer of cuticule (Doweld

2000), n = 12, size strongly bimodal.

Usually included in the Balanophorales, but differ

from them markedly in monocarpellate gynoecium

with conduplicate stylodium and deﬁ ned ovarian cav-

ity, not reduced ovule, presence of stomata, pollen

grains with reticulate ornamentation, and presence of

bisexual ﬂ owers.

1. CYNOMORIACEAE

Endlicher ex Lindley 1833. 1/2. From the Canary

Islands, the Mediterranean coastal regions, from

Morocco to Egypt and western Asia to Mongolia.

Cynomorium.

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Order 104. BALANOPHORALES

Terrestrial herbs with yellowish-white to yellow,

orange to red, or brown colors parasiting roots of trees,

shrubs, or rarely herbs. Attaching to the roots of the

host by an often irregularly lobed or branched, some-

times more or less horizontal and elongated tuberous

organ that ranges from several centimeters to the size

of a human head. Vascular system very reduced, but

sometimes contains vessels with simple or (?) scalari-

form perforations. Sieve-element plastids of S-type.

Stems with or without leaves, unbranched. Leaves

much reduced (but subterranean only), or absent.

Leaves when present, membranous, scaly, usually

spiral, rarely opposite, decussate or distichous, very

rarely verticillate (Balanophora involullata), without

stomata. Inﬂ orescences terminal, racemose, spicate or

spadix-like; branches subtended by scaly caducous

bracts, sometimes with sterile apical part peltately

widened. Flowers usually very small (those of some

genera among the smallest in ﬂ owering plants), numer-

ous or very numerous, unisexual (monoecious or

dioecious), apetalous and frequently without calyx,

entomophilous. Male ﬂ owers often with 2(3) (4–8)

free or basally connate, valvate sepals. Stamens usu-

ally 3–4 or more opposite to the sepals or less often

only 1–2 stamens (1 stamen in Dactylanthus and

2 stamens in Mystropetalon and Lophophytoideae),

free and with tetrasporangiate anthers or more or less

united into a synandrium; synandria opening irregu-

larly; anthers extrorse, usually connate, opening longi-

tudinally or (Balanophora subgenus Balania)

transversally. Tapetum secretory. Microsporogenesis

simultaneous (successive – Corynaea?). Pollen grains

2-celled or 3-celled (Mistropetaloideae, Helosidoideae),

3- colpate, rugate, 3–6-colporate, 3–5-porate or

3(7–8)-pororate or rarely inaperturate (Balanophora

subgenus Balania), smooth or variously sculptured.

Female ﬂ owers commonly with the calyx tube com-

pletely fused with the ovary wall, or the calyx appar-

ently absent, but in Mystropetalon there is still 3-lobed

upper free part of the calyx, which in some other

genera is 2-lobed or irregularly lobed or reduced to an

Superorder BALANOPHORANAE 427

inconspicuous, lobed rim or represented by a few cells

at the top of the ovary. Gynoecium of three or two

completely fused carpels with more or less distinct

stylodia (Helosidoideae and Lophophytoideae), stylo-

dia united into a 1–3-lobed style (Mystropetaloideae,

Dactylanthoideae, and Balanophoroideae) or stigma

sessile (Sarcophytoideae). Ovary superior to inferior,

1–3-locular (Mystropetalon) or more often unilocular,

with one ovule per locule, which is sometimes erect

on the ﬂ oor of the cavity or more commonly suspended

from the roof by a stalklike cell or (Langsdorfﬁ a and

Thonningia) completely united with the wall of the

ovary. Ovules ategmic, with reduced nucellus, usually

little more than a female gametophyte. In Balanophora

even the placenta is completely reduced. Female

gametophyte bisporic of Allium-type or monosporic of

Polygonum-type. Endosperm cellular, with large cha-

lazal haustorium. Fruits 1-seeded, indehiscent, often

very tiny, nutlets or drupelets, in Mystropetalon

surrounded by the swollen calyx tube and with dis-

clike, whitish elaiosome (modiﬁ ed pedicel) at the

base. Seeds with a minute, undifferentiated embryo

surrounded by copious endosperm; cotyledons lack-

ing; testa probably always absent. Contain starch or a

waxy resinous substance, balanophorin (Langsdorfﬁ a

and Balanophora). n = 14, 16, 18.

Both Cynomoreales and Balanophorales are proba-

bly related to the Santalales.

1. BALANOPHORACEAE

A. Richard 1822 (including Dactylanthaceae Takhtajan,

1987, Hachetteaceae Doweld 2001, Helosidaceae

Bromhead 1840, Langsdorfﬁ aceae van Tieghem ex

Pilger et K. Krause 1914, Lophophytaceae Bromhead

1840, Mystropetalaceae J. D. Hooker 1853,

Sarcophytaceae A. Kerner 1891, Scybaliaceae

A. Kerner 1891). 16/50. Tropical and South

(Mystropetalon) Africa, Madagascar, Comoros,

Tropical Asia, Malesia, Paciﬁ c Islands, tropical

Australia, New Caledonia (Hachettea), New Zealand

(Dactylanthus), Tropical America.

1.1 MYSTROPETALOIDEAE

Cells with abundant deposits of reddish-brown

substance (mystrin) present in leaves, superﬁ cial layers

of the cortex of axis, interspersed among the vascular

tissue and present also in bracts, ﬂ owers, and fruits.

Calyx of the male ﬂ owers 3-merous and zygomorphic,

gamosepalous. Stamens two, ﬁ laments adnate to lower,

narrow part of the adaxial sepals. Calyx of the female

ﬂ owers extremely small, cuplike, slightly zygomor-

phic. Ovary inferior, completely adnate to the calyx

tube, 3-merous, basally encircled by an irregularly

crenate disc; from the base of the calyx cup arises the

relatively long ﬁ liform style, which ends in a papillate,

more or less clearly 3-lobed capitate stigma. Ovules

three, pendulous. – Mystropetalon.

1.2 DACTYLANTHOIDEAE

Male ﬂ owers with or without petals. Stamens 1–2, free

or united. Female ﬂ owers with a small tubular, 3-lobed

calyx or with 2 ﬁ liform sepals. Stylodia united into a

style. Pollen grains pantoporate. Seed coat lacking. –

Hachettea, Dactylanthus.

1.3 SARCOPHYTOIDEAE

Inﬂ orescence conspicuously branched, paniculate,

with ﬂ eshy axis, bracteate (Sarcophyte) or

(Chlamydophytum) ebracteate or minutely bracteate.

Stamens 3–4 or 7–10, free. Pollen grains 3-porate.

Stigma sessile. Ovary with 3–1 ovules. – Sarcophyte,

Chlamydophytum.

1.4 SCYBALIOIDEAE

Stem with triangular, scaly leaves. Inﬂ orescence when

young covered by triangular, imbricate bracts. Pollen

grains mostly 2-celled, pantoporate. – Scybalium.

1.5 HELOSIDOIDEAE

Stem without scaly leaves. Inﬂ orescence when young

covered by hexagonal, marginally coherent peltate

bracts. Pollen grains 3-celled, 3-colpate. – Helosis

(including Latraeophila and Exorhopala

), Corynaea,

Ditepalanthus, Rhopalocnemis.

1.6 LOPHOPHYTOIDEAE

Both female and male ﬂ owers mostly naked.

Stylodia distinct, 2. Pollen grains 3-colporate.

Flowers in clavate panicles, not immersed in a layer of

chaffy hairs. – Lophophytum, Lathrophytum,

Ombrophytum (including Juelia).

1.7 LANGSDORFFOIDEAE

Female ﬂ owers with tubular irregularly lobed or

dentate calyx. The ovule is completely united with the

428 Subclass VI. ROSIDAE

wall of the ovary. The stigmatoid tissue of the style is

conspicuously enveloped. – Langsdorfﬁ a, Thonningia.

1.8 BALANOPHOROIDEAE

Female ﬂ owers without calyx. The ovule is pendulous

in the cavity of the ovary. The stigmatic tissue is not

morphologically distinguished from the rest of the

style. – Balanophora (including Balania?).

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Superorder RHAMNANAE 429

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Superorder RHAMNANAE

Order 105. RHAMNALES

Trees and shrubs, rarely woody climbers, or subshrubs,

very seldom herbs. Branches, leaves and ﬂ owers often

covered with simple (mostly), stellate or peltate

trichomes. Vessels mostly with simple perforations,

rarely scalariform to reticulate; lateral pitting alternate.

Fibers with simple or bordered pits. Rays from mark-

edly heterogeneous to homogeneous. Axial paren-

chyma predominantly paratracheal in most species,

predominantly apotracheal, sometimes very scanty.

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

trilacunar. Leaves alternate or opposite, simple,

pinnately veined or with several main veins from the

base, with small stipules or estipulate. Stomata mostly

anomocytic, less often paracytic or anisocytic. Flowers

in various kinds of terminal or axillary inﬂ orescences,

sometimes solitary, bisexual or rarely unisexual (plants

dioecious), actinomorphic, 5–6-merous or more rarely

3–4-merous, sometimes apetalous. Sepals 5–6, or 3–4,

usually basally connate, valvate, deciduous or seldom

persistent. Petals 3–6, rarely 8, often inserted on the

calyx tube, sometimes wanting (Elaeagnaceae).

Stamens 4–9(-12), free, alternisepalous; ﬁ laments thin

or very short (Barbeyaceae and Elaeagnaceae), free or

adnate to petals; anthers generally 2-locular, basiﬁ xed

or dorsiﬁ xed, introrse, latrorse or extrorse, tetrasporan-

giate, opening longitudinally. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains

2-celled or 3-celled, usually 3-colporate, often more or

less triangular. Nectary disc usually well developed.

Gynoecium of (1-)2–3(-5) united carpels, rarely

(Dirachmaceae) carpels 8; style long with cylindrical,

linear or capitate stigmas; ovary superior to inferior,

1-locular or rarely 5–6(-8)-locular. Ovules solitary,

rarely 2 (in Karwinskia), anatropous, with inwardly

directed micropyle, bitegmic, crassinucellate. Female

gametophyte of Polygonum- or Allium-type. Endosperm

nuclear. Fruits drupaceous, dry, and indehiscent or

eventually dehiscent, or separating into dehiscent

or indehiscent mericarps, baccate, rarely samaroid, or

capsular, or achenes. Seeds exotestal, arillate or more

often exarillate; mesotesta with a few sclerotic cells,

embryo usually straight, endosperm starchy, usually

scanty, sometimes wanting. Usually producing glyco-

sides, often with various kinds of alkaloids, ﬂ avonols,

n = (6–8)10–14(23).

Probably related to the Celastrales, from which they

are chieﬂ y distinguished by the antepetalous stamens.

Both Rhamnales and Celastrales evidently had a com-

mon origin from a diplostemonous rosanean stock, the

antepetalous or antesepalous cycles having aborted.

Key to Families

1 Flowers with developed nectaries. Alkaloids

present.

2 Gynoecium of 2–3(-5) united carpels. Trees and

shrubs, often lianoid (by aid of hooks in Ventilago,

tendrils in Gouania, twining stems in Berchemia),

or xerophytic (sometimes with ﬂ attened, spiny,

assimilatory stems and reduced leaves, such as

Colletia armata), rarely (Crumenaria) subshrubs

or even (Crumenaria decumbens) herbs. Tendrils

climbering or scrambling. Hairs mostly simple,

430 Subclass VI. ROSIDAE

stellate in Pomaderreae. Some members of the

family, such as ssp. of Ceanothus and Colletia,

contain nitrogen-ﬁ xing actynomycets in the

roots. Mucilage cells are common in the leaf and

in the primary cortex of the axis, and mucilage

cavities are sometimes present as well in the lat-

ter. Vessels mostly with simple perforations,

rarely scalariform to reticulate; lateral pitting

alternate. Fibers with simple pits. Rays from

markedly heterogeneous to homogeneous. Axial

parenchyma predominantly paratracheal in most

species, predominantly apotracheal in a few spe-

cies. Nodes trilacunar. Leaves alternate or oppo-

site, simple, pinnately veined or with several

main veins from the base, sometimes much

reduced, with very small stipules (sometimes

modiﬁ ed into spines), wanting in most species of

Phylica. Stomata mostly anomocytic, less often

paracytic or anisocytic. Flowers mostly rather

small, greenish, in various kinds of terminal or

axillary inﬂ orescences (sometimes reduced to a

solitary ﬂ ower), bisexual or rarely unisexual

(plants dioecious), actinomorphic, hypanthium

present, 5–6-merous or more rarely 4-merous,

with more or less cuplike ﬂ oral tube that is often

circumscissile and deciduous above the middle.

Calyx 5–4-lobed, lobes triangular, valvate and

are individually deciduous or seldom persistent.

Petals usually smaller than sepals and more or

less concave or hooded, and holding anthers fre-

quently clawed at the base, rarely wanting.

Stamens 5–4, free, alternisepalous; ﬁ laments

thin, adnate to base of petals; anthers minute,

dorsiﬁ xed, introrse, generally 2-locular. Pollen

grains 2-celled usually 3-colporate, often more

or less triangular. Nectary disc usually well

developed, intrastaminal, usually adnate to the

ﬂ oral tube and sometimes to the ovary. Gynoecium

of 2–3(-5) carpels, with a terminal lobed or often

deeply cleft style; in 3-carpellate ovaries the

sequence is usually fertile-sterile-fertile: one

septum bears 2 ovules, the second – a single

ovule, and the third is sterile (Medan and

Schirarend 2004); ovary superior to inferior, or

fully or imperfectly multilocular (rarely

pseudomonomerous), with 1 (2 in Karwinskia)

basal and erect ovule in each locule. Ovules with

inwardly directed micropyle. Fruits drupaceous,

dry, and indehiscent or eventually dehiscent, or

separating into dehiscent or indehiscent meri-

carps, baccate, rarely samaroid or capsular. Seeds

exotestal, sometimes with dorsal groove, arillate,

sometimes with a large aril (Alphitonia) or more

often exarillate; mesotesta with a few sclerotic

cells, endotegmen of cuboid cells, with scalari-

form thickenings; embryo large, green, oily, usu-

ally straight, rarely (Reynosia) ruminate;

endosperm starchy, usually rather scanty (rumi-

nate in Reynosia), sometimes wanting. Usually

producing anthraquinone glycosides, often with

various kinds of alkaloids (sometimes benzyliso-

quinoline alkaloids), ﬂ avonols, n = (6–8)10–

13(23).. . . . . . . . . . . . . . . . . . . 1. rhamnaceae.

2 Gynoecium of 1 carpel. Shrubs or small trees,

rarely woody climbers; shoots often reduced to

spines; cambium storied. Leaves, young branches,

and calyx tube are covered with peltate and stel-

late trichomes; mycorrhiza present in the cortex of

the rootlets of Hippophae and Shepherdia, and

nodules on the roots of all three genera contain

nitrogen-ﬁ xing bacteria. Vessels with simple per-

forations; lateral pitting alternate, rather small.

Fibers with conspicuously bordered pits. Rays

weakly heterogeneous to homogeneous in some

species of Elaeagnus. Axial parenchyma diffuse,

sometimes very scanty. Phloem usually stratiﬁ ed

tangentially into hard and soft layers. Nodes uni-

lacunar. Leaves alternate or (Shepherdia) oppo-

site, simple, entire, pinnately veined, estipulate.

Stomata anomocytic. Flowers in racemose inﬂ o-

rescences or sometimes solitary in the axils of

leaves, bisexual (spp. of Elaeagnus) or more

rarely polygamous (spp. of Elaeagnus) or dioe-

cious (Shepherdia and Hippophae), actinomor-

phic, mostly 4-merous, apetalous. Calyx often

somewhat corolloid, tubular to saucer-shaped,

usually 4-lobed or very rarely 6-lobed, in

Hippophae with only 2 lobes, valvate. Stamens

borne in the throat of the calyx tube, equal in

number and alternating with the sepals

(Elaeagnus), or twice as many and both alternate

with and opposite to them (4 in Hippophae and 8

in Shepherdia), ﬁ laments very short; anthers tet-

rasporangiate, dorsiﬁ xed or basiﬁ xed. Pollen

grains 2-celled or 3-celled, (2)3(4)-colporate.

More or less well-developed, often lobed nectary

disc usually present on the inner side of the calyx

tube. Gynoecium of 1 carpel (probably as a result

Superorder RHAMNANAE 431

of complete abortion of the other carpels), with an

elongate, slender stylodium ending in a linear or

capitate stigma; ovary superior, 1-locular. Ovule

solitary, basal, with a short and broad funicle, with

a funicular obturator. Female gametophyte of

Polygonum- or (Shepherdia) Allium-type.

Endospermal chalazal haustorium develops in

Elaeagnus. Fruits achenes with thin, membranous

pericarp enveloped by the persistent base of the

calyx tube that becomes mealy or ﬂ eshy, often

with a long, inner layer. Seeds have a hard testa

and contain a straight embryo with large, thick,

plano-convex cotyledons and a short radicle point-

ing downward; endosperm scanty around the

embryo (Shepherdia), only near the radicle

(Hippophae), or wanting (Elaeagnus). Strongly

tanniniferous, usually producing quebrachitol and

sometimes also indole alkaloids, sinapinic and

ellagic acids, ﬂ avonols, 0-methyl ﬂ avonoids, trit-

erpenes, n = 6, 10, 11, 13, 14. 2. elaeagnaceae.

1 Flowers without nectaries. Alkaloids absent.

3 Leaves alternate. Deciduous small trees or

shrubs. Young annual shoots and leaves with

unicellular hairs. Leaves clustered on the short

shoots, rapidly covered with long unicellular

trichomes on the abaxial side, simple, dentate-

serrate, with persistent subulate stipules.

Stomata anomocytic or cyclocytic. Flowers

solitary and axillary, pedicellate, bisexual, acti-

nomorphic, with an epicalyx of 4–8 bracteoles.

Hypanthium present between the petals and the

calyx lobes. Sepals 5–6 or 8, basally connate,

externally densely pubescent and in Dirachma

somalensis also with glandular peltate hairs (as

well as on pedicels), valvate. Petals 5–6 or 8,

free, inserted on the calyx tube, imbricate, with

ﬂ eshy and densely haired appendages near base

that are covering a separate cavity in which

nectar is exuded by a ﬂ at epithelial gland or

nectar is secreted by the basal part of the

appendages. Stamens 5–6 or 8, opposite the

petals, free, inserted on calyx tube above the

petals; ﬁ laments subulate, in D. somalensis

with two diverging short spurs at the lower ven-

tral part, propping the petal appendages; anthers

large, 2- locular, oblong-ellipsoidal, basiﬁ xed,

latrorse or extrorse. Pollen grains 3-colporate,

reticulate. Gynoecium of 8 carpels; style long,

with cylindrical or linear stigmas; ovary supe-

rior, tomentose, 5–6- or 8-locular; ovule soli-

tary in each locule, ascending from the inner

angle, anatropous, hypotropous, bitegmic;

micropyle zig-zag. Fruits septicidal and septi-

fragal, 8-locular (Dirachma socotrana) or

5-locular (D. somalensis), woolly inside with

very long, unicellular trichomes and separating

from base to apex into ventrally dehiscent seg-

ments. Seeds ellipsoidal, with straight embryo

and scanty endosperm; seed coat with exotesta

and endotegmic pigment layer. Contain ﬂ a-

vonoids. . . . . . . . . . . . . . . . 4. dirachmaceae.

3 Leaves opposite. Small trees; the wood with-

out growth rings and very hard. Vessel ele-

ments very short, with oblique end walls

provided by simple perforations; lateral pit-

ting alternate. Fibers with simple or bordered

pits. Rays somewhat heterogeneous, mixed

uniseriate and pluriseriate. Axial parenchyma

very scanty, diffuse. Sieve-elements plastids

of S-type, with very oblique, compound sieve

plate. Nodes unilacunar, with a single trace.

Leaves petiolate, decussate, simple, entire,

pinnately veined, densely white-tomentose

below with curled, unicellular hairs, estipu-

late. Stomata paracytic. Flowers small, in short

3-ﬂ owered dichasia, dioecious, actinomor-

phic, apetalous, without bracts and bracteoles.

Sepals 3 or 4, slightly connate at the base, val-

vate; those of the female ﬂ owers pinnately

veined and slightly imbricate, accrescent in

fruit. Stamens 6–9 (up to 12), with very short,

free ﬁ laments; anthers elongate, apiculate,

latrorse. Pollen grains 3-colporate, tectate-

columellate, with granular layer beneath the

thick, perforated tecturn. Exine ornamentation

rugulate with spinules, colpi with thickened

margins, and pores covered with exinous

islets. Gynoecium of 1 (mostly) to 3 free or

basally more or less connate carpels, each with

linear stylodium; stigma linear, papillous all

round; ovary superior, sessile to shortly stipi-

tate, 1-locular; ovules solitary, pendulous,

subapical ovule on a short funiculus, anatro-

pous, bitegmic (Bouman and Boesewinkel

1997), crassinucellate. Endosperm develop-

ment nuclear. Fruits dry, indehiscent, nut-like,

with thin pericarp and accrescent, submem-

branous, prominently veined sepals. Seeds

432 Subclass VI. ROSIDAE

unspecialized, remain parenchymatic and

locally compressed, exotesta perforated,

endotegmen tanniniferous; embryo straight,

endosperm scanty; cotyledons ﬂ at, ﬂ eshy.

Embryo and endosperm contain aleuron

grains. Accumulating ellagic acid and ﬂ avonols

quercetin and kaempferol. . 3. barbeyaceae.

1. RHAMNACEAE

A.L. de Jussieu 1789 (including Frangulaceae A.P. de

Candolle 1805, Gouaniaceae Raﬁ nesque 1837,

Phylicaceae J.G.Agardh 1858, Ziziphaceae Adanson

ex Post et Kuntze 1903). 52/900. Nearly cosmopolitan,

but mainly in tropical and subtropical regions.

Classiﬁ cation after D. Medan and C. Schirarend

(2004).

PALIUREAE: Paliurus, Ziziphus (incl. Sarcomp-

halus), Hovenia; colletieae: Trevoa (incl. Talguenea),

Retanilla, Adolphia, Kentrothamnus, Colletia,

Discaria; phyliceae: Phylica, Trichocephalus,

Nesiota, Noltea; gouanieae: Gouania, Helinus,

Reissekia, Alvimi antha, Crumenaria, Pleuranthodes;

pomaderreae: Pomaderris, Siegfriedia, Trymalium,

Spyridium, Stenanthemum, Cryptandra, Blackallia;

rhamneae: Rhamnus (including Frangula), Scutia,

Sageretia, Berchemiella, Rhamnella (including

Chaydaia), Dallachya, Berchemia, Rhamnidium,

Karwinskia, Condalia, Auerodendron, Reynosia,

Krugiodendron; maesopsideae: Maesopsis; venti-

lagineae: Smythea, Ventilago; ampelozizipheae:

Ampelozi ziphus; doerpfeldieae: Doerpfeldia;

bathior hamneae: Bathiorhamnus.

Incertae sedis: Alphitonia, Ceanothus, Colubrina,

Emmenosperma, Granitites, Lasiodiscus, Schistocarpaea.

2. ELAEAGNACEAE

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

Meyer 1836). 3/50. Subtropical and temperate regions

of the Northern Hemisphere and in tropical Asia, with

1 species of Elaeagnus in Queensland; Shepherdia (3)

is endemic to North America.

Elaeagnus, Hippophae, Shepherdia.

Share many features with the Rhamnaceae, inclu-

ding seed anatomy (“The seed-structure is typically

Rhamnaceous,” states Corner [1976:124]). The afﬁ nity

with the Rhamnaceae is also conﬁ rmed by uredino-

logical data (see Holm 1979 and Savile 1979).

A common origin is very probable.

3. BARBEYACEAE

Rendle 1916. 1/1. Northeastern Africa and adjacent

parts of the Arabian Peninsula.

Barbeya.

Have many similarities with Rhamnaceae, includ-

ing wood anatomy and ﬂ ower morphology.

4. DIRACHMACEAE

Hutchinson 1959. 1/2. Socotra and central Somalia.

Dirachma.

The genus Dirachma is usually included in the

Ger-aniales with some members of which it has certain

similarities, such as beaked fruits dehiscing from

below to apex. However, Hutchinson (1959, 1967,

1968, 1973) included his family into Tiliales. Link

(1991, 1993), Ronse Decraene and Smets (1995) have

found Dirachma more closely related to the Malvales

than to any other group, including Geraniales. They

especially emphasize the location and anatomy of epi-

thelial nectaries, which in some important details

resemble trichomatic nectaries of Grewieae, Coloneae,

Duboscieae, Lueheeae, Trichospermeae, and

Triumpheteae of the Tiliaceae as well as the develop-

ment of long, unicellular trichomes on the fruit valves’

ventral sides known in some Bombacaceae, such as

Ceiba and Ochroma. Also, Dirachma has mucilage

cells (Yakovleva 1994), stellate hairs, hypocalyx, hap-

lostemonous androecium and hypotropous ovules

(epitropous in Geraniaceae). However, Thulin et al.

(1998) conclude that Dirachmaceae are closely related

to the Barbeyaceae. But Dirachma differs from

Barbeya in many importance characters, including

alternate and stipulate leaves, bisexual ﬂ owers, 8–9

sepals not accrescent in fruit, pollen morphology, the

number of carpels, bitegmic ovules, septicidal 5–8-loc-

ular capsules, seed anatomy and also in wood structure

such as the thicker vessel walls, smaller vessel pits,

scanty or entirely lacking axial parenchyma, absence

of vasicentric or vascular tracheids, and lack of pris-

matic crystals in the wood. Boesewinkel and Bouman

(1997) proposed relationships Dirachmaceae to the