Takhtajan Armen. Flowering Plants

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Superoder CORNANAE 463

Shaw 1965) and as well as in not branched aperture oper-

culum elements (Backlund and Nilsson 1997). I there-

fore to accept the Triplostegiaceae as a separate family

very closely related to the Valerianaceae.

7. DIPSACACEAE

A.L. de Jussieu 1789 (including Scabiosaceae

Martynov 1820). 14/300. Temperate Eurasia (espe-

cially in Mediterranean region and western Asia) and

tropical Africa and South Africa.

Cephalaria, Dipsacus, Succisa, Succisella, Knautia,

Ptero cephalus, Pterocephalidium, Scabiosa, Lom-

elosia, Pseudoscabiosa, Scabiosiopsis, Sixalix, Tremas-

telma, Pycnocomon.

Probably originated from some Triplostegia-like

ancestor whose open inﬂ orescence condensed into a

ca-pitulum (see Burtt 1977: 106).

8. MORINACEAE

Raﬁ nesque 1820. 3/13. From the Balkans (Greece,

Bulgaria, Romania) and Lebanon and Israel to conti-

nental China.

Morina, Acanthocalyx, Cryptothladia

The distinctness of the family Morinaceae is con-

ﬁ rmed by palynological and embryological data

(Vinokurova 1959; Vijayaraghavan and Sarveshwari

1968; Verlaque 1977; Kamelina 1980; Blackmore and

Cannon 1983) as well as by morphology of the inﬂ o-

rescence and basic chromosome numbers. Morinaceae

pollen is morphologically distinct from that of the

Dipsacaceae in the lack of columellae and operculae

and in the presence of zonorate endoapertures in some

of the pollen types (Blackmore and Cannon 1983).

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Order 113. AQUIFOLIALES

Trees and shrubs, sometimes woody lianas. Vessels

with scalariform or (Icacinaceae) scalariform and sim-

ple perforations, or simple perforation; lateral pitting

scalariform to alternate. Fibers usually with distinctly

bordered pits. Rays heterogeneous, with elongate or

short ends. Axial parenchyma apotracheal to paratra-

cheal. Sieve-element plastids of S-type. Nodes trilacu-

nar or less often unilacunar. Leaves alternate or rarely

opposite, simple, estipulate or (most of Aquifoliaceae

and Helwingiaceae) with very small and caduceus

stipules. Stomata of diverse types. Flowers small,

greenish or whitish, in cymose or racemose inﬂ ores-

cences or rarely solitary, bisexual or unisexual, actino-

morphic, usually 4-merous or 5-merous. Sepals free or

more or less connate. Petals free or less often connate,

imbricate or valvate, rarely wanting. Androecium

haplostemonous with antesepalous stamens; anthers

tetrasporangiate, usually opening longitudinally.

Tapetum secretory. Microsporogenesis simultaneous.

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

Gynoecium mostly of 2–5 united carpels, with short

style or sessile stigma. Ovary superior or inferior, with

1–2 pendulous ovules in each locule. Ovules com-

monly anatropous, unitegmic or (Sphenostemonaceae

and Cardiopteridaceae) bitegmic, crassinucellate to

tenuinucellate, sometimes (Aquifoliaceae) with endo-

thelium. Female gametophyte of Polygonum-type.

Endosperm cellular (Aquifoliaceae and Nothapodytes

Superoder CORNANAE 467

in Icacinaceae) or nuclear (Gonocaryum in Icacinaceae)

(the other families not studied for nuclear development).

Fruits drupaceous, baccate, or samara. Seeds exotestal,

with very small to large embryo; endosperm present or

absent. Icacinaceae contain iridoid compounds.

According to molecular date (Olmstead et al. 2000;

Soltis et al. 2000, 2006; Bremer et al. 2002) Aquifoliales

related to Dipsacales and Apiales.

Key to Families

1 Gynoecium of 3–5 carpels.

2 Ovary superior.

3 All (2)4–6 locules of the ovary fertile.

Evergreen or deciduous trees (usually small)

or shrubs, sometimes climbers and epiphytic;

resiniferous and laticiferous idioblasts pres-

ent. Vessels with exclusively scalariform per-

foration, and without vestured pits, usually

with numerous bars; lateral pitting usually

opposite, rarely scalariform or alternate. Fibers

often septate. Rays heterogeneous with elon-

gate ends. Axial parenchyma diffuse. Nodes

unilacunar, or trilacunar. Leaves alternate or

very rarely opposite, or whorled, entire, leath-

ery, toothed or spinose, often with resiniferous

or laticiferous cells in mesophyll; stipules very

small and often caducous or wanting. Stomata

mostly anomocytic. Flowers very small, in

axillary or supra axillary fascicles or cymes,

dioecious or occasionally polygamous or

(Nemopanthus) polygamo-dioecious, 4-mer-

ous or less often 5-merous, rarely 6–9-merous.

Sepals small, more or less connate at the base,

imbricate, in Nemopanthus minute and decid-

uous in male ﬂ owers and wanting from female

ones. Petals free (Nemopanthus) or shortly

connate at base (Ilex), imbricate. Stamens usu-

ally as many as and alternate with the petals,

mostly adnate to the petals at base, sometimes

more numerous than the petals in polycarpous

species of Ilex. Staminodia and pistillodia

present. Pollen grains 3-colporate. Gynoecium

usually of 4–5 carpels, rarely less (2–3) or

more (up to 24); style short or nearly wanting,

with lobed or capitate stigma; ovary with soli-

tary or very rarely two ovules in each locule;

ovules anatropous, unitegmic, crassinucellate

to tenuinucellate (two species of Ilex), with

endothelium. Fruits drupes with as many

pyrenes as carpels. Seeds with cuboid exot-

estal cells, tangentially elongated, inner walls

ligniﬁ ed, rest crushed, endotesta tanninifer-

ous; embryo near micropyle, very small;

endosperm copious, oily and proteinaceous,

without starch. Plants sometimes producing

alkaloids and ﬂ avonols (kaempferol and quer-

cetin), n = 9, 10. . . . . . . . .1. aquifoliaceae.

3 Fertile locule solitary, rarely all three locules

fertile. Trees and shrubs, sometimes lianas

with non- axillary branch tendrils, laticiferous,

or non-laticiferous and without coloured juice.

Trichomes unicellular. Stem often with anom-

alous structure, sometimes with interxylary

phloem. Vessels scalariform, scalariform and

simple, or (in stems with unilacunar nodes)

only simple perforations; vessels without ves-

tured pits; lateral pitting usually intermediate

between scalariform and opposite or (in ves-

sels with simple perforations) alternate. Fibers

usually with bordered pits. Rays heteroge-

neous with elongate or short ends. Axial

parenchyma apotracheal or less often paratra-

cheal. Nodes trilacunar or less often unilacu-

nar. Leaves alternate or less often opposite

(Iodes), leathery, petiolate, entire to toothed,

estipulate. Stomata anomocytic, anisocytic, or

paracytic. Inﬂ orescences axillary, sometimes

terminal, or opposite leaves (Citronella),

racemes, thyrsoids, cymes, pseudospikes (pis-

tilate plants of Calatola). Flowers small,

bisexual or less often polygamous to dioe-

cious, (3)4–5(6)-merous. Sepals more or less

connate, imbricate or rarely valvate, rarely

wanting. Petals free or rarely basally connate,

usually valvate, rarely wanting. Stamens as

many as and alternate petals; ﬁ laments free or

borne on corolla tube alternate with its

lobes; anthers usually introrse, dorsiﬁ xed,

opening longitudinally or less often by api-

cal pores. Pollen grains 2-celled, aperturate,

or nonaperturate (Stachyanthus), colpate, or

porate, or colporate, or foraminate. Nectary

disc mostly absent, but sometimes present or

represented by distinct glands. Gynoecium of

(2)3(-5) carpels with short style; stigma punc-

tiform, subcapitate, capitate, or lobed; ovary

usually 1-locular by abortion of the remaining

locules; rarely (Emmotum) 3-locular; ovules

468 Subclass VII. ASTERIDAE

two, rarely one per locule, pendulous, anatro-

pous, unitegmic but with free integuments at

the micropyle in Phytocrene, crassinucellate

to tenuinucellate; funicle usually thickened

near the micropyle. Fruits usually drupaceous,

often ﬂ attened and or ribbed, 1-seeded, rarely

dry and winged. Seeds with straight or curved,

green embryo; endosperm mostly well devel-

oped, oily, sometimes wanting. Plants some-

times with iridoid compounds, ﬂ avonols

(quercetin); alkaloids and proanthocyanidins

present or absent; verbascosides present

(Cassinopsis), n = 10, 12. . . . 2. icacinaceae.

2 Ovary inferior. Glabrous dioecious shrubs. Vessels

with scalariform perforations that have 17–41

widely spaced bars; lateral pitting scalariform.

Fibers with minutely bordered pits in radial walls,

all septate. Axial parenchyma scanty paratracheal.

Rays heterogeneous. Silica grains present in ray

cells, and axial parenchyma, and also in septate

ﬁ bres of Helwingia himalaica. Nodes unilacunar.

Leaves alternate to subopposite, spiral, petiolate,

simple, remotely serrate or crenate-serrate, ovate

to lanceolate, with deciduous stipules often

divided into ﬁ liform lobes. Stomata anomocytic.

Flowers unisexual, in small umbels mostly borne

on the leaves through the axillary peduncle being

adnate to the upper side of midrib, the males sev-

eral in an umbel, the female 1–3; bracts absent,

male ﬂ owers sometimes with more or less long

pedicel, female ﬂ owers with very short pedicels.

Calyx limb obsolete; petals 3–5, valvate, continu-

ous with the calyx tube. Stamens 3–5, alternipe-

talous, inserted outside ﬂ at angled nectary disc;

anthers short. Pollen grains 3-colporate, with

perforate tectum and diffuse endoaperture.

Gynoecium of 3–4 carpels; ﬂ at nectary disc

forming a cone passing into short style with 3–4

recurved stigmas; ovary inferior, 3–4-locular, with

one pendulous, anatropous, with dorsal raphe,

unitegmic, tenuinucellate ovule per locule. Fruits

ovoid or globose drupes, angular when dry, with

3–4 separate cartilaginous or crustaceous pyrenes.

Seeds with minute and nearly globose embryo at

the apex of copious endosperm; seed-coat thin,

consists of 2–3 layers compressed, thin-walled

cells. Contain caffeic acid (chlorogeneic acid),

n = 19.. . . . . . . . . . . . . . . . . . 3. helwingiaceae.

1 Gynoecium of two carpels.

4 Stipules present. Trees or shrubs, glabrous, ter-

restrial or epiphytic; young stem with separate

bundles. Vessels with scalariform perforations;

lateral pitting opposite. Fibers with bordered pits.

Rays heterogeneous. Axial parenchyma apotra-

cheal. Leaves alternate, entire or serrate, acumi-

nate; stipules small, ﬁ mbriate, caduceus, with

glandular hairs on adaxial surface. Inﬂ orescences

epiphyllous, ﬂ owers borne on the leaf toward the

tip of the midrib. Flowers very small, actinomor-

phic, bisexual, green or white, fasciculate

(cymose false umbels), 5-merous. Sepals con-

nate into obconic tube adnate to ovary, open,

toothed, persistent. Petals inserted below the

margins of the epigynous ﬂ at nectary disc,

spreading, ovate-triangular, acute, valvate.

Stamens ﬁ ve, with short, subulate ﬁ laments and

small, bilobed anthers. Pollen grains 3-colporate,

with complete tectum and granules. Gynoecium

of two carpels and two small, recurved stigmas

on very short stylodia; ovary inferior, incom-

pletely 2-locular, with usually numerous 2-seri-

ate ovules on two parietal placentas. Fruits small,

incompletely 2-locular, 3–6-seeded berries.

Seeds oblong, slightly curved, rugose; testa mul-

tilayered, exotestal cells large, thick-walled,

mucilaginous; embryo very minute at the base of

ﬂ eshy endosperm. . . . . . . 4. phyllonomaceae.

4 Leaves estipulate.

5 Ovary 2-locular. Evergreen trees and shrubs.

Vessels with scalariform perforations that have

32–185 bars; lateral pitting scalariform; phloem

stratiﬁ ed. Leaves alternate or subopposite or

pseudo-verticillate, simple, leathery, glandular-

dentate to entire. Flowers in subterminal, rac-

emose inﬂ orescences, bracteate, bisexual,

actinomorphic, 4-merous. Sepals free, imbri-

cate, caducous. Petals free, ﬂ eshy, caducous, or

wanting. Stamens 4–6, or 8–12, ﬁ laments short

and thick or obsolete; anthers with thick con-

nective, elongate and latrorse or short and

introrse, petaloid or ﬁ lantherous. Stamens lam-

inal with embedded microsporangia. Pollen

grains 3-perorate, reticulate to scabrate.

Gynoecium of two carpels with large sessile

capitate stigma. Ovary superior, 2-locular, with

1(2), pendulous ovules in each carpel; ovules

anatropous, bitegmic, crassinucellate. Fruits

drupelike (berry, according to Bailey 1956,

Superoder CORNANAE 469

pseudo-drupe – van Steenis 1986), with sepa-

rable pyrenes or with one stone (with 1–2 bony

pyrenes), usually more or less asymmetrical,

ovoid, crowned by the stigma (Watson and

Dallwits 2006). Pericarp ﬂ eshy, ﬁ nally black.

Seeds more or less ruminate, exotestal or exo-

endotestal, endotestal cells with dark contents;

embryo small, surrounded by a thick, chalazal

envelope, of which the outer layer is hard and

bony and whether or not ruminate functioning

as and superﬁ cially resembling the structure of

a pyrene (van Steenis 1986), endosperm copi-

ous, oily. . . . . . . . . . 5. sphenostemonaceae.

5 Ovary 1-locular. Climbing herbs with articu-

lated laticifers bearing abundant milky juice in

the leaves and stems. Vessel elements short and

very broad, with simple perforations; lateral

pitting alternate. Fibers with small bordered

pits. Nodes trilacunar with three traces. Leaves

alternate, thinly membranous, simple or lobed

to varying degrees, long-petioled, cordate,

3–7-veined from the base, estipulate. Stomata

anomocytic and paracytic. Flowers very small,

in axillary branched, more or less scorpioid

cymes, bisexual or polygamous, actinomorphic,

4–5-merous. Calyx (4-)5-lobed, lobes imbri-

cate, persistent. Corolla shortly sympetalous,

(4-)5-lobed, lobes spreading, thin, imbricate,

caduceus. Stamens (4-)5, inserted midway

down the corolla tube, or in the throat of the

corolla tube and alternate with the sepals; ﬁ la-

ments very short, glabrous; anthers introrse,

dorsiﬁ xed, tetrasporangiate, opening longitu-

dinally. Pollen grains 3-colporate, with striate

ornamentation. Nectary disc wanting. Gynoecium

of two united carpels; stylodia two, dissimilar,

one longer, thicker, cylindrical or subclavate,

persistent on fruit, the other very short, thin-

ner, with capitate stigma; ovary superior, 1-loc-

ular, with two ovules pendulous from the

summit of the locule, one of which often abor-

tive. Ovules anatropous, bitegmic, crassinucel-

lete, with dorsal raphe. Fruits obovate-oblong,

dry, indehiscent, ﬂ at, 1-seeded, broadly

2-winged samara, more or less stipitate; wings

transversely striolate. Seeds linear, sulcate,

exotestal, embryo minute, well differentiated,

at the apex of very densely granular; endosperm

ﬂ eshy. . . . . . . . . . . . . . 6. cardiopteridaceae.

1. AQUIFOLIACEAE

A. Richard 1828 (including Ilicaceae Berchtold et

J. Presl 1820). 2/400–450. Ilex is very widely distrib-

uted; Nemopanthus (2) is found only in eastern North

America.

Ilex (including Byronia), Nemopanthus

One is the most archaic families which is charac-

terized by primitive wood structure, the absence of

nectarines, and very small embryo.

2. ICACINACEAE

Mires 1851. 53/400 (including Emmotaceae Van Tieghem

1899; Iodaceae Van Tieghem 1899; Leptaulaceae

Van Tieghem 1899; Metteniusaceae H. Karsten ex

Schnizlein 1843–1870; Pennantiaceae J.G. Agardh

1858; Phytocrenaceae Arnott ex R. Browne 1852;

Pleurisanthaceae Van Tieghem 1899; Sarcostigmataceae

Van Tieghem ec Bullock 1958; Stemonuraceae Kårehed

2001). Pantropical, with relatively few species in tem-

perate regions.

icacineae: Citronella (including Villaresia),

Emmotum, Poraqueiba, Oecopetalum, Pittosporopsis,

Pseudobotrys, Platea, Gonocaryum, Medusanthera,

Ottoschulazia, Gomphandra, Rhyticaryum, Hartleya,

Codiocarpus, Medusanthera, Apodytes, Casimirella,

Leretia, Pleurisanthes, Merrilliodendron, Icacina,

Mappia, Lasianthera, Cantleya, Nothapodytes,

Stemonurus, Whitmorea, etc.; discophoreae:

Discophora; iodeae: Mappianthus, Polyporandra,

Iodes, Hosiea, Natsiatum, Natsiatopsis; phytocreneae:

Pyrenacantha (including Trematosperma), Miquelia,

Phytocrene, etc.; sarcostigmateae: Sarcostigma;

METTENIUSEAE

: metteniusa.

Insertae sedis: Alsodeiopsis, Chlamydocarya,

Desmostachys, Lavigeria, Polycephalium, Stachyanthus,

Cassinopsis, Calatola, Raphiostylis, Dendrobangia,

Leptaulus, Gastrolepis, Grisollea, Irvingbaileya.

Share many common features with the Aquifoliaceae.

3. HELWINGIACEAE

Decaisne 1836. 1/5. Eastern Himalayas (from central

Nepal to Bhutan), northeastern India (Assam and

Mantpur), northern Burma, continental China, northern

Vietnam, Taiwan, Ryukyu Islands, Japan.

470 Subclass VII. ASTERIDAE

Helwingia

Helwingiaceae related to Aquifoliaceae and

Phyllonomaceae (Soltis et al. 2006).

4. PHYLLONOMACEAE

Small 1905 (Dulongiaceae J. Agardh 1858, nom. illeg).

1/4. Mexico to northwestern Bolivia.

Phyllonoma (Dulongia)

According to Soltis and Soltis (1997), “Analyses

or rbcL sequences similarly revealed that Phyllo-

noma is allied with traditional asterids, with a

particu larly close relationship between Phyllonoma

and Helwingia”.

5. SPHENOSTEMONACEAE

P. Royen and Airy Shaw 1972. 1/7–10. Central Malesia

to New Guinea, tropical Australia, and New Caledonia.

Sphenostemon

Close to the Aquifoliaceae and especially to

Icacinaceae (Baas 1975).

6. CARDIOPTERIDACEAE

Blume 1847. 1/3 (including Peripterygiaceae G. King

1895). Northeastern India (Assam and Bengal), Burma,

Southeast Asia, Malesia from Malay Peninsula to New

Guinea, and Queensland.

Cardiopteris (Peripterygium)

Related to the Icacinaceae, but differ in bitegmic

ovules and fruit type.

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472 Subclass VII. ASTERIDAE

Order 114. APIALES

Trees, shrubs, woody lianas, or herbs, glabrous or with

various types of trichomes. Mostly with schizogenous

secretory canals or cavities containing ethereal oils,

resins, and gums usually well developed in various

parts, especially in the parenchymatous tissues. Pith

bundles and cortical bundles often present. Vessels

with simple or sometimes with scalariform, or simple

and scalariform perforations; lateral pitting scalari-

form to alternate. Fibers with simple or (Toricelliaceae)

bordered pits, often septate. Rays heterogeneous

to seldom homogeneous. Axial parenchyma apo-

tra cheal (Toricelliaceae, Myodocarpus, Delarbrea,

Porospermum, Pseudosciadium) or more often paratra-

cheal. Sieve-element plastids of S-type. Nodes multila-

cunar, sometimes pentalacunar, or rarely trilacunar.

Leaves alternate or rarely opposite or verticillate, sim-

ple to compound; petioles usually broad and sheathing,

sometimes stipulate, commonly with a ring or arc of

vascular bundles, and sometimes with medullary bun-

dles as well; leaf mesophyll cells contain oil bodies.

Stomata paracytic, anomocytic, or of other types.

Flowers usually small, in terminal or axillary umbels,

that are very often arranged into secondary umbels or

sometimes into other types of compound inﬂ ores-

cences, rarely in racemes or spikes, bisexual or seldom

unisexual, actinomorphic or the outermost ones some-

times more or less zygomorphic, mostly 5-merous.

Calyx usually represented by small teeth around the

top of the ovary, which often much reduced or even

wanting. Petals nearly always free, valvate or some-

times (some Araliaceae) more or less imbricate, rarely

wanting. Stamens as many as and alternate with the

petals, sometimes twice as many as petals or even

(some Araliaceae) numerous; anthers mostly tetraspo-

rangiate, opening longitudinally. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains

3-celled or rarely 2-celled, mostly 3-colporate or

(Toricelliaceae) 4-colporate. Gynoecium of 2–5(-many)

united carpels; stylodia free or more or less connate

into a style that is often enlarged at the base into a sty-

lopodium conﬂ uent with the nectary disc; ovary infe-

rior, or semi-inferior, rarely superior, crowned by the

glandular nectary disc, with one pendulous ovule per

carpel (but an abortive ovule sometimes present).

Ovules anatropous, pendulous, epitropous, with ven-

tral or dorsal raphe and with micropyle directed upward

and outward, unitegmic, crassinucellate to tenuinucel-

late, usually with endothelium. Female gametophyte of

Polygonum-type or sometimes of Oenothera-, Allium-,

Penaea-, or Drusa-types. Endosperm nuclear. Fruits

drupaceous or baccate or more often schizocarps con-

sisting of two indehiscent, 1-seeded mericarps usually

separating in the plane of union of two carpels, before

falling the two mericarps usually hang for a time on a

commonly forked stalk, the carpophore, which devel-

ops from the carpellary tissues. Seeds exotestal, with

small (sometimes minute), straight embryo and copi-

ous endosperm. Usually containing sesquiterpenes,

triterpenoid compounds, caffeic acid derivatives, and

polyacetolenes, iridoids and ellagic acid present or

lacking. Seeds containing petroselinic acid.

Related to both the Hydrangeales and Cornales, espe-

cially to the former. Probably had a common origin.

Key to Families

1 Ovary superior.

2 Secretory canals absent.

3 Petiole without sheathing base. Shrubs or

trees. Trichomes unicellular, recurved or mul-

ticellular, uniseriate. Vessesl with scalariform

perforation and with bordered pits. Rays het-

erogenous. Axial parenchyma apotracheal or

scanty paratracheal. Fibers nonseptate. Nodes

trilacunar with three traces. Leaves alternate,

simple, serrate or entire, petiolate, glabrous,

estipulate. Stomata paracytic. Flowers in ter-

minal panicles, with articulated pedicels, uni-

sexual (plants dioecious or polygamous),

5-merous. Calyx minute, imbricate, caducous.

Petals glabrous, free, valvate. Stamens ﬁ ve,

free; ﬁ laments ﬁ liform, connate; anthers

2-locular, introrse, dorsiﬁ xed. Pollen grains

3-colporate. Nectary disc absent. Gynoecium

of two carpels. Ovary superior, 1-locular,

without ﬂ eshy appendage; stigma 3-lobed, on

a very short style, or discoid and sessile. Ovule

solitary and even though a second abortive

may be present, pendulous. Fruit small drupe,

stone obtusely trigonous; seed coat thin;

embryo short, minute, endosperm present.

Leuco-anthocyanins and caffeic acid present;

n = 25.. . . . . . . . . . . . . . . 1. pennantiaceae.

3 Petioles with sheathing base. Woody plants.

Vessels with simple or almost exclusively

scalariform perforations, without helical