Takhtajan Armen. Flowering Plants

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Superoder CORNANAE 473

thickenings. Axial parenchyma apotracheal.

Rays homogeneous, with simple or bordered

pits. Leaves alternate, pinnately compound or

simple, entire, estipulate. Inﬂ orescences pan-

icles of umbellules. Flowers with articulated

pedicels, unisexual or bisexual, 5-merous.

Sepals large, imbricate. Petals free, imbricate,

caducous. Nectary disc present. Gynoecium

of two carpels. Ovary inferior, 2-locular.

Fruits ﬂ eshy or spongy drupes (Delarbrea),

or dry schizocarps (Myodocarpus); seeds

laterally ﬂ attened; woody endocarp hard,

with large oil ducts; endosperm smooth and

has druses, in Myodocarpus up to 18 druses

in cell (Vyshenskaya 2000).; n = 12.

. . . . . . . . . . . . . . . . . . . 8. myodocarpaceae.

2 Secretory canals present. Small trees, shrubs,

often woody lianas, sometimes subshrubs, gla-

brous or with simple, uniseriate trichomes with

two or three basal cells and a long terminal cell,

two-armed hairs with a short stalk, or club-

shaped glandular hairs. A constant and charac-

teristic feature is the presence of well-developed,

schizogenous secretory canals in the pericycle of

the stem, roots, and leaves, and also in the sec-

ondary cortex and secondary phloem of older

stems. Vessels with simple perforations or very

rarely scalariform perforations (in primary xylem

always with scalariform perforations); lateral pit-

ting alternate, small to very small. Fibers septate,

with small, simple, or very narrowly bordered

pits. Rays nearly homo geneous. Axial paren-

chyma scanty paratracheal. Sieve-element plas-

tids of Ss-type. Nodes trilacunar, rarely

unilacunar. Leaves alternate, sometimes very

closely crowded at the branch tips, simple, entire,

or rarely serrate, sinuate or occasionally more or

less lobed (but seedling leaves and juvenile

leaves may be pinnatiﬁ d), sometimes with wavy

margins, coriaceous, estipulate. Stomata para-

cytic. Flowers in corymbs or cymose panicles or

solitary, bisexual or seldom poly gamous (in sev-

eral New Zealand spp. of Pittosporum), or uni-

sexual, actinomorphic or rarely (Cheiranthera)

slightly zygomorphic, 5-merous (except for

gyno ecium), bibracteolate. Sepals imbricate, free

or some times basally connate. Petals usually

basally connate forming more or less distinct

tube, with imbricate lobes. Stamens alternate

with petals, free or weakly connate basally;

anthers tetrasporangiate, opening longitudinally

or rarely (Cheiranthera) by apical pores. Nectary

disc absent. Microsporogenesis simultaneous.

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

Gynoecium of two or rarely (spp. of Pittosporum)

3–5 carpels, with simple style and capitate or

slightly lobed stigma; ovary superior, 1-locular,

with parietal placentas or as a result of protrusion

and fusion the placentas become 2- or rarely 3-loc-

ular, with several to usually more or less numerous

ovules per placenta or per locule. Ovules anatro-

pous or nearly campylotropous, tenuinucellate.

Female gametophyte of Polygonum-type. Fruits

loculicidal (sometimes also septicidal) capsules

(Pittosporeae) or berries (Billardiereae). Seeds

often immersed in a viscid pulp, with a minute

embryo placed near the hilum; endosperm copi-

ous, oily, and proteinaceous; seed coat 6–12

cells thick, unspecialized. Lack iridoid com-

pounds and ellagic and gallic acid but contain

ethereal oils, resins and mucilages, leuco-

anthocyanins, caffeic, quinic, and sinapic acids,

ﬂ avonols kaempferol and quercetin, polyacety-

lenes, and often triterpenoid saponins, n = 12.

. . . . . . . . . . . . . . . . . . . . . . . 6. pittosporaceae.

1 Ovary usually inferior.

4 Fruits nearly always carpels, drupaceous, or bac-

cate. Trees, shrubs, woody lianas, or woody epi-

phytes, rarely perennial herbs; plants bearing

essential oils, or without essential oils. Vessels soli-

tary (Pentapanax, Arthrophyllum, Myodocarpus,

Delarbrea, Boerlagio dendron, Oreopanax,

Plerandra, Dendropanax, some species of

Schefﬂ era and Poly scias – Oskolski 1996), or in

short radial multiples, mostly with simple perfora-

tions, in a few genera with scalariform perfora-

tions (Myodocarpus exclusively scalariform) that

have few to numerous bars (up to 56 in

Boerlagiodendron); lateral pitting scalariform to

alternate. Fibers with small, simple pits, often

septate, but in Boerlagiodendron ﬁ ber tracheids.

Rays mostly heterogeneous with short ends or

rarely homogeneous. Axial parenchyma usually

scanty paratracheal, rarely (Myodocarpus,

Delarbrea) diffuse-in-aggregates or (Reynoldia)

metatracheal. Nodes multilacunar, pentalacunar,

or seldom trilacunar. Leaves alternate or very

rarely (Cheirodendron) opposite or verticillate,

474 Subclass VII. ASTERIDAE

simple, dissected, pinnately or palmately com-

pound, sometimes to second or third degree, aro-

matic or not, gland-dotted or not; petiole usually

broad and sheathing, stipulate or estipulate;

stipules when present, intrapetiolar, often adnate

to and scarcely distinguishable from the base of

the petiole. Stomata mostly paracytic or aniso-

cytic, rarely anomocytic. Flowers usually small,

mostly in simple umbels, bisexual or rarely polyg-

amous or dioecious, rarely calyptrate, usually

more or less 5-merous. Calyx teeth small, often

much reduced or even obsolete. Petals 5–10(-12),

rarely three, free or rarely connate at the base or

forming a calyptra (as in Tupidanthus) or in a tube

(as in Boerlagiodendron), deciduous, valvate or

sometimes slightly imbricate. Stamens usually as

many as and alternate with petals or rarely twice

as many or numerous (up to 120 in Tupidanthus),

anthers dorsiﬁ xed, tetrasporangiate or sometimes

of as many as eight microsporangia. Pollen grains

3-celled, usually 3-colporate, sometimes 2- or

4-colporate, commonly reticulate. Gynoecium of

2–5 to many (up to 200 in Tupidanthus), mostly

ﬁ ve carpels; stylodia free or sometimes more or

less connate into a style, generally swollen at the

base to form a more or less deﬁ nite stylopodium

that is conﬂ uent with the epigynous nectary disc;

ovary inferior or semi- inferior, very rarely supe-

rior, 1-locular (Seemannaralia) or pseudo-

monomerous, with one (sometimes the second

abortive one) pendulous ovule per locule; ovules

crassinucellate to tenuinucellate. Female gameto-

phyte usually monosporic, of Polygonum-type,

Fruits drupes with as many pyrenes as carpels, or

berries, rarely a drupelike schizocarp, which in

Stilbocarpa and Astrotricha, has carpophores,

Porospermum have oily glands in endocarp,

Mackinlaya has glands in pericarp. Seeds with

small embryo and copious, ruminate or not, oily

endosperm containing large amount of petrose-

linic acid. Endosperm of Tetraplasandra, Arthro-

phyllum, and Aralia has druses (Vyshenskaya

2000). Alkaloids and ﬂ avonols (quercetin and

kaempferol) present or absent; n = 11, 12 + (usu-

ally) or more. . . . . . . . . . . . . . . . .7. araliaceae.

4 Fruits usually schizocarps with persistent carpo-

phore. Mostly perennial or less often annual

herbs, sometimes subshrubs, rarely shrubs or

soft-wooded small trees. Plants with essential

oils, or without, resinous, or not. Stems usually

with well-developed collenchyma and large pith

that shrinks or dries at maturity. Internodes com-

monly becoming hollow. Secretory cavities with

resin, oil, or with mucilage. Vessels almost

always with simple perforations, or rarely with

scalariform, very rarely with vestigial bars.

Nodes multilacunar. Leaves small to large, usu-

ally alternate, or alternate and opposite (the upper

sometimes more or less opposite), usually much

dissected (ternate or variously pinnate) infre-

quently entire, and sometimes more or less paral-

lelveined, gland-dotted or not, aromatic or foetid,

or rarely without marked odour, estipulate, but

sometimes with stipular ﬂ anges; petioles usually

with broad sheathing base. Stomata anomocytic

or paracytic. Flowers small, mostly compound

umbels or less often in heads or simple umbels,

rarely (as in Petagnia) in dichasial inﬂ orescence;

at the top of the stalk of each individual umbel

(umbellule) an involucre of bracts is often found

(the bracts of outer ﬂ owers), and a similar large

involucre often occurs at the top of the main stalk

bearing the compound umbel. Flowers mostly

bisexual or seldom unisexual and monoecious

(as Echinophora) or dioecious (as Acronema and

Arctopus) actinomorphic or often peripheral

ﬂ owers more or less zygomorphic, usually 5-mer-

ous except for 2-merous gynoecium. Calyx teeth

usually more or less reduced and frequently

inconspicuous, but in Saniculeae they are well

developed. Petals free, typically inﬂ exed at the

tip, valvate, rarely wanting. Stamens ﬁ ve, alter-

nate with the petals, borne on the epigynous

nectary disc; anthers dorsiﬁ xed or basiﬁ xed,

introrse, tetrasporangiate; ﬁ laments long; anthers

introrse. Pollen grains 3-celled, 3-colporate, very

rarely 2-colporate, with more or less smooth sur-

face. Gynoecium of two rather reduced carpels

with free, short stylodia usually set on variously

and characteristically developed swollen base

(stylopodia); ovary inferior, 2-locular or rarely

1-locular and pseudomonomerous, with one ovule

per carpel (sometimes with an additional abortive

ovule); ovules anatropous, tenuinucellate or

pseudocrassinucellate. Female gametophyte

mostly of Polygonum-type. Fruits non-ﬂ eshy

schizocarps; mericarps two, dry, united facially,

1-seeded, the integument sometimes united with

Superoder CORNANAE 475

the pericarp, always separating at maturity. Seeds

adherent to the pericarp, with usually small, well

differentiated, straight embryo and copious, oily

endosperm containing large amount of petrose-

linic acid. Alkaloids present (and usually toxic

via falcorinone) or absent; ﬂ avonols (kaempferol

and quercetin) present or absent; n = 4–12, most

commonly 8 or 11. . . . . . . . . . . . . . 9. apiaceae.

5 Stomata anomocytic. Small trees or shrubs,

glabrous. Secretory cavities absent. Hairs sim-

ple, glandular. Vessels mostly with simple per-

forations with distinct rims or occasionally

scalariform, with 5–24 bars. Fibers with

minutely bordered pits, some ﬁ bers septate.

Rays heterogeneous. Axial parenchyma scanty

apotracheal. Crystals absent (Noshiro et Baas

1998). Leaves alternate, entire or dentate,

long-petiolate, petioles more or less sheathing

at base, stipulate or have shortly stalked glan-

dular hairs with spherical heads, blackening

on drying. Stomata amonocytic. Flowers in

axillary or terminal racemes or panicles,

bisexual, 5-merous; pedicels bracteate at the

base and mostly 2-bracteolate at the apex.

Sepals connate into a minutely 5-dentate tube,

imbricate. Petals free, imbricate, reﬂ exed.

Stamens ﬁ ve; ﬁ laments short, slender; anthers

oblongate, basiﬁ xed or dorsiﬁ xed. Pollen

grains 3-colporate. Nectary disc obscure or

obsolete. Gynoecium of 2–3-carpels; stylodia

2–3, free, subulate, erect or recurved, with lin-

ear or punctiform stigma; ovary inferior,

2–3-locular (only one of which is fertile), with

one pendulous, anatropous ovule per locule.

Fruits ovoid or oblong drupaceous, with one

fertile dorsal and two sterile ventral loculi, the

septa marked by external grooves. Seeds light

brown, with very thin testa; embryo minute,

straight, differentiated; endosperm ﬂ eshy,

copious.. . . . . . . . . . 5. melanophyllaceae.

5 Stomata encyclocytic. Trees or shrubs, often

epiphytic or climbing. Secretory canals absent.

Hairs all unicellular, provided with granular,

nodular, or verrucose prominences. Wood dif-

fuse-porous. Vessels with scalariform perfora-

tions of 10–17 (up to 28) bars. Fibers with

distinctly bordered pits on both radial and tan-

gential walls. Rays heterogeneous. Axial

parenchyma diffuse-in-aggregates. Nodes

penta- or trilacunar. Prismatic crystals usually

in body ray cells. Leaves alternate, often asym-

metrical, coriaceous, simple, entire or angled

or spinose-dentate, estipulate; epidermal cells

rhomboidal, small; palisade cells entire, cylin-

dric; hypodermis present; petioles dilated and

slightly sheathing at base, with arc of bundles.

Flowers small, in terminal or axillary panicles

or racemes, dioecious, pedicels ebracteolate,

articulated. Male ﬂ owers: calyx minute, 5-den-

tate; petals ﬁ ve, imbricate; stamens ﬁ ve, alter-

nipetalous; anthers broadly oblong,

tetrasporangiate, dorsiﬁ xed, introrse, opening

longitudinally. Pollen grains 3-colporate.

Nectary disc ﬂ eshy, 5-sided. Pistillodia lack-

ing. Female ﬂ owers: calyx tube ovoid or tur-

binate, 5-dentate; petals imbricate or absent;

gynoecium of three united carpels; stylodia

three, very short, free, and/or shortly connate,

stigmatose at the apex or inside; ovary infe-

rior, 1–2-locular (only one of which is fertile),

with one pendulous ovule in each locule.

Staminodia lacking. Ovules anatropous, uni-

tegmic, crassinucellate, with endothelium.

Female gametophyte of Polygonum-type.

Fruits ovoid, usually 1-locular and 1-seeded

berries. Seeds oblong, with small, elongate,

straight embryo in copious endosperm.

Contain iridoid glucoside griselinoside and

caffeic acid (chlorogenic acid), but lack gallic

and ellagic acids. Seeds contain petroselinic

acid, n = 18. . . . . . . . . . . 2. griselianaceae.

6 Funiculus thickened to form an obturator.

Small trees with thick branches and broad

pith. Secretory canals absent. Hairs both

simple unicellular (in ﬂ oral parts sometimes

with transverse wall) and characteristic

multicellular glandular hairs consisting of

three or four cells. Primary cork with typi-

cal collenchyma. Vessel elements short,

with simple perforations; lateral pitting

opposite to alternate, without vestures pits.

Fibers with minutely bordered pits, septate.

Rays heterogeneous with long ends. Axial

parenchyma scanty paratracheal. Nodes

multilacunar. Crystals absent. Leaves alter-

nate, simple, more or less palmatilobed,

entire or more often coarsely toothed, pal-

mately veined, long-petiolate, broadly

476 Subclass VII. ASTERIDAE

sheathed at base, estipulate. Stomata anomo-

cytic. Flowers small, in lax, multiﬂ owered,

axillary panicles (thyrses), dioecious, actino-

morphic, bracteolate. Male ﬂ owers: pedicel

articulate; calyx 5-lobed, with short tube,

lobes open, more or less unequal; petals

ﬁ ve, induplicate-valvate, concave, apex

elongate, inﬂ exed; stamens ﬁ ve, alternipeta-

lous; ﬁ laments short; anthers tetrasporan-

giate, basiﬁ xed, opening longitudinally,

latero-introrse; pollen grains 4-colporate,

with short colpi, simple endoaperture and

either granulose or superﬁ cially rugulate

tectum with processes or projections that are

«short capillilike» and in section are den-

ticulate; nectary disc more or less ﬂ at; pistil-

lodia of 1–3 subulate processes in the middle

of disc. Female ﬂ owers: pedicel not articu-

late; calyx minutely and irregularly

3–5-toothed; petals wanting; staminodia

absent; nectary disc inconspicuous or absent;

gynoecium of 3–4 carpels, with three short,

thick, subulate, erect, or divaricate, papil-

lose free stylodia; ovary inferior, 3–4-locu-

lar with one apical, pendulous ovule per

locule (often partly sterile); ovules anatro-

pous, apotropous, with micropyle directed

inward. Fruits obliquely ovoid 3–4 locular,

usually 1-seeded, rarely 2-seeded drupe

crowned by persistent stylodia and with

more or less circular opening into each

empty chamber. Seeds linear with small,

curved embryo at the apex of copious, ﬂ eshy

endosperm. Contains iridoid compound

griselinoside; n = 12. . .4. toricelliaceae.

6 Funiculus not thickened. Small trees. Hairs

both unicellular and strongly asymmetrical,

glandular trichomes. Secretory canals

absent. Wood diffuse- porous. Pith with

groups of sclereids. Vessels with scalari-

form perforations with up to 34 bars; lateral

pitting opposite or sometimes scalariform.

Fibers with minutely bordered pits mostly

in radial walls, all septate; walls thin to

thick. Rays multiseriate (up to 12 cells

wide), heterogeneous. Axial parenchyma

mostly paratracheal. Nodes multilacunar.

Crystals absent. Leaves large, alternate,

entire or more often deeply and irregularly

pinnately lobed or partite with an end lobe,

estipulate; epidermal cells large, sinuate,

palisade cells lobed, tubular; petioles dilated

and sheathing at base, with scattered bun-

dles. Stomata anisocytic. Flowers small

(buds c 2.5 mm long), creamy or red-tinged,

in large terminal much branched panicles or

cymules, dioecious, actinomorphic. Pedicels

articulated. Calyx with ﬁ ve very small teeth.

Petals ﬁ ve, imbricate. Stamens ﬁ ve, alterni-

petalous; ﬁ laments ﬂ attened, anthers round,

tetrasporangiate, dorsiﬁ xed, opening longi-

tudinally. Pollen grains 3-colporate, tectate,

smooth, minutely perforate. Stamens in

female ﬂ owers sterile (staminodia). Nectary

disc intrastaminal, represented in male

ﬂ owers by uniform cushion of tissue and in

female ﬂ owers by three (rarely four) gib-

bous stylopodia. Gynoecium of three (rarely

four) carpels; stylodia 3(-4), free, short,

diverging, subulate from a broad base (sty-

lopodium), with terminal stigma; ovary

inferior, with three (rarely four) loculi, of

which all but one remain small with no ovu-

lar rudiments; the fertile locule with one

pendulous ovule. Ovule anatropous, with

dorsal raphe, unitegmic (with very massive

integument), tenuinucellate. Fruits drupa-

ceous, large, ellipsoid, exocarp with net-

work surface and inconspicuous grooves

near the apex, which represent incompletely

ligniﬁ ed vestiges of empty locules; meso-

carp ﬂ eshy; endocarp hard, ridged. Seeds

large, with 4–5 grooves; endosperm deeply

ruminate, enclosing small embryo at the

micropylar end. Plants contain the iridoid

glucosides griselinoside and aralioside, and

caffeic acid. n = 20. . . . .3. aralidiaceae.

1. PENNANTIACEAE

J.G. Agardh 1858. 1/4. Eastern Australia (1, P. cun-

ninghamii), New Zealand (3).

Pennantia

The genus Pennantia has been assigned by different

authors to Rhamnaceae, Euphorbiaceae, Terebintha-

ceae, Anacardiaceae, Icacinaceae. Considering these

uncertainties, Agardh (1858) placed this genus in a

Superoder CORNANAE 477

family of its own. According to recent studies

Pennantiaceae belong to the Apiales (Kårehed 2001,

2003).

2. GRISELINIACEAE

J.R. Forster et G. Forster ex A. Cunningham 1839. 1/7.

New Zealand, Chile, Argentina, Paraguay, south-

eastern Brazil.

Griselinia

Usually Griselinia is included in the Cornaceae (in

the tribe Griselinieae Wangerin, which contains two

genera – Griselinia and Melanophylla). However,

as Philipson (1967) and others have demonstrated,

Griselinia differs from Cornaceae in a number of

important characters, including alternate leaves,

nodal anatomy, the presence of a hypodermis below

the upper epidermis of leaves, encyclocytic stomata,

the presence of spiral thickenings in both vessels

and ﬁ bers, imbricate petals, free or only shortly con-

nate stylodia, usually 1-locular ovary, and baccate

fruits. Both in wood anatomy (Metcaife and Chalk

1951) and ﬂ oral vasculature (Philipson 1967) the

Griseliniaceae have some similarities with Araliaceae.

Molecular data also supported transfer Griselinia into

Apiales.

3. ARALIDIACEAE

Philipson et B.C. Stone 1980. 1/1. Southern Thailand,

peninsular Malaysia, Sumatra, Singapore, the Anambas

Islands, and Borneo.

Aralidium

The genus Aralidium has been assigned by different

authors to both the Araliaceae and the Cornaceae.

From the Cornaceae it differs mainly in multilacunar

nodes, pinnatiﬁ d leaves, leaf sheaths, endocarp mor-

phology, pollen morphology, and ruminate endosperm.

From the Araliaceae it differs in the absence of secre-

tory canals, peripheral origin of the ventral bundles of

the ovary (they rise through the ovary wall and not the

axis, and the central axis of the gynoecium has no vas-

cular bundles), the dorsal position of the raphe, the

absence of the rudiment of second ovule, the presence

of glandular hairs, and anisocytic stomata. In spite of

some features it has in common with the Araliaceae,

including sheathing base of the petiole, and some fea-

tures of wood anatomy, the genus Aralidium is not

closely related with the Araliaceae and related fami-

lies. With the latter Aralidium shares characters such

as dioecious ﬂ owers, multilacunar nodes, paratracheal

axial parenchyma, functionally unilocular ovary, three

free stylodia, and the presence of griselinoside

(Philipson et al. 1980).

4. TORICELLIACEAE

H. Hu 1934. 1/3. Eastern Himalayas (from western

Nepal to Bhutan), northern Burma, continental west-

ern China.

Toricellia

The genus Toricellia is usually included in the

Cornaceae. However, it differs from them in many

characters, including multicellular glandular hairs,

vessels with simple perforations, the presence of typi-

cal collenchyma, free stylodia, the presence of obtura-

tor, and especially in its very distinctive pollen type

(Ferguson and Hideux 1978). Most probably related to

the Aralidiaceae (Eyde 1988: 313).

5. MELANOPHYLLACEAE

Takhtajan ex Airy Shaw 1972. 1/6–8. Madagascar.

Melanophylla

Diverse opinions have been expressed as to the

afﬁ nities of this remarkable Madagascan genus.

Generally, it is included in Cornaceae. However, the

genus Melanophylla differs markedly from Cornaceae

in alternate leaves with sheathing petioles, glandular

hairs with spherical heads, racemose inﬂ orescences,

imbricate petals, and free stylodia. Plankett (2001) and

Lundberg both suggest that the Melanophyllaceae

and Griseliniaceae are successive lineages near the

base of Apiales. Chandler and Plunkett (2004) include

Melanophylla in their “Torricellia group”.

6. PITTOSPORACEAE

R. Brown 1814. 12/c.240. Tropical and subtropical

regions of the Old World, mostly Australia (nine

endemic genera); the largest and the only widely dis-

tributed genus Pittosporum (ca. 200) occurs from

southern Africa to New Zealand and the Paciﬁ c Islands

478 Subclass VII. ASTERIDAE

with one outlier, P. coriaceum, on the island of Madeira

and in a single locality in Tenerife.

pittosporeae: Pittosporum, Auranticarpa,

Rhytido sporum, Bursaria, Hymenosporum; billar-

diereae: Citriobatus, Pronaya, Billardiera,

Marianthus, Bentl eya, Sollya, Cheiranthera.

Pittosporaceae have many features in common with

the Araliaceae and most probably originated from the

same protoaralealean stock. In some respects

Pittosporaceae are even less specialized than Araliaceae

(usually well-developed calyx and superior ovary), but

in some other respects, including wood anatomy, they

are more advanced. As early as 1884, van Tieghem

suggested that Pittosporaceae are probably rather

closely related to the Araliaceae and Apiaceae. He

especially emphasized the possession of schizogenous

secretory canals and the similarity in the origin of lat-

eral roots in both families (secretory canals situated

externally to the xylem inhibit the development of lat-

eral roots on the outer side of the xylem groups, and

consequently they arise between the secretory passage

opposite a xylem group and the neighboring secretory

passage opposite a phloem group; see Metcaife and

Chalk [1950: 129]). Later Solereder (1899) and Guenot

(1906) have suggested on anatomical grounds that

Pittosporaceae are related to the Araliaceae and

Apiaceae. Schurhoff (1929) studied the embryology of

Pittisporum and came to the same conclusion. Huber

(1963) arrived at a similar conclusion. Finally,

Hegnauer (1969) concluded that on phytochemical

grounds the family is unlike members of the saxifraga-

ceous stock (accepted by many authors) and that the

inclusion of the family in the Araliales «appears much

more likely to reﬂ ect its natural incorporation in the

Resales.» According to Jay (1969), the Pittosporaceae

should be placed near the Apiaceae rather than near the

Saxifragaceae.

7. ARALIACEAE

A.L. de Jussieu 1789 (including Botryodendraceae

J.G. Agardh 1858; Hederaceae Giseke 1792; Hydro-

cotylaceae N. Hylander 1945). 47–52/1325–1350.

Tropical and subtropical regions, with relatively few

genera in temperate regions; concentrated mostly in

southern and southeastern Asia and Paciﬁ c islands

(especially New Caledonia); one of the most archaic

genus Myodocarpus is endemic to New Caledonia.

7.1 HYDROCOTYLOIDEAE

Mostly perennial herbs. Nodes trilacunar with three

traces. Leaves rounded-peltate, crenate; stipules cau-

line. Stomata anisocytic. Gynoecium of two carpels.

Carpophore undivided. – Hydrocotyle, Trachymene,

Neosciadium.

7.2 ARALIOIDEAE

Trees, shrubs, woody lianad, rarely herbaceous plants.

Nodes usually multilacunar or pentalacunar. Leaves

pinnately or palmately compound, stipulate or estipu-

late, stipules when present intrapetiolar. Stomata

mostly paracytic or anisocytic, rarely anomocytic.

Gynoecium mostly of ﬁ ve carpels. Carpophore usually

absent. – schefflereae: Schefﬂ era (including

Plerandra), Tupidanthus, Schefﬂ eropsis, Didymo-

panax, Eleutherococcus (Acanthopanax), Kalopanax,

Opio panax, Tetrapanax, Merilliopanax, Dendropanax,

Boerlagiodendron, Osmoxylon, Trevesia, Fatsia,

Gastonia, Cussonia, Reynoldsia, Tetraplasandra,

Munroidendron; meryteae: Meryta; hedereae:

Hedera, Gamblea, Oreopanax, Astrotricha, Trevesia,

Kissodendron, Brassaiopsis, Macropanax, Hetero-

panax, Arthrophyllum, Wardenia; aralieae:

Polyscias, Pseudopanax, Raukaua, Sciadodendron,

Pentapanax, Motherwellia, Sciadodendron, Aralia,

Cephalaralia, etc; panaceae: Harmsiopanax,

Panax. ?? Anakasia, Cheirodendrom.

8. MYODOCARPACEAE

Doweld 2001. 2/17. Eastern Malesia, New Caledonia,

Vanuatu, Australia (Queensland).

Delarbrea (including Porospermum,

Pseudosciadium), Myodocarpus.

Related to the Pittosporaceae and Araliaceae, but

closely related to the Araliaceae.

9. APIACEAE

Lindley 1836 or Umbelliferae A.L. de Jussieu 1789

(nom. altern.) (including Actinotaceae Konstantinova

et Melikian 2006; Ammiaceae Barnhard 1895; Angeli-

caceae Martynov 1820; Bupleuraceae Martynov 1820;

Coriandraceae Burnett 1835; Daucaceae Martynov

1820; Eryngiaceae Raﬁ nesque 1838; Ferulaceae

Saccardo 1872; Imperatoriaceae Martynov 1820;

Superoder CORNANAE 479

Mackinlayaceae Doweld 2001; Saniculaceae A. Löve

et Löve 1974; Smyrniaceae Burnett 1835). 435/3100–

3200. Cosmopolitan, but mainly temperate regions of

the Northern Hemisphere (particularly Mediterranean

and Irano-Turanian regions).

Closely related to the Araliaceae.

9.1 APIOIDEAE

Fruits with a soft parenchymatous endocarp, some-

times hardened by woody subepidermal layers.

Stylodia on apex of disc. – echinophoreae: Echino-

phora, Anisosciadium, Dicyclophora, Pycnocycia,

Thecocarpus, Ergocarpon; scandiceae: Scandix, Physo-

caulis, Chaerophyllum, Grammosciadium, Anthriscus,

Myrrhis, Geocaryum, Rhabdosciadium, Myrrhoides,

Osmorhiza, Sphallerocarpus, Kozlovia, Krasnovia,

Balansaea, Molopospermum, Tinguarra, Noecono-

podium, Athamanta, Conopodium, Oreomyrrhis,

Ptychotis, Hladnikia; heteromorpheae: Hetero-

morpha, Anginon, Dracosciadium, Glia, Polemannia;

caucalideae: Astrodaucus, Caucalis, Daucus, Exoa-

cantha, Lisaea, Orlaya, Torilis, Turgenia, etc.;

coriandreae: Schrenkia, Coriandrum, Bifora, Koso-

pol janskia, Fuernrohria, etc.; smyrnieae: Scaligeria,

Smyrnium, Oreomyrrhis, Smyrniopsis, Conium,

Tauschia, Arracacia, Lecokia, Trachydium, Cachrys,

Prangos; PLEUROSPERMEAE: Pleurospermum,

Eleutherospermum, Aulacospermum, Physospermum;

ERIGENIEAE: Erigenia; hohenackerieae: Hohena-

ckeria; pyramidoptereae: Pyramidoptera; apieae:

Bupleurum, Trinia, Szovitsia, Aphanopleura, Froriepia,

Apium, Petroselinum, Cicufa, Trachyspermum, Crypto-

taenia, Ammi, Falcaria, Carum, Bunium, Chamaes-

ciadium, Conopodium, Pimpinella, Aegopodium. Slum,

Crithmum, Seseli, Oenanthe, Aethusa, Athamanta,

Foeniculum, Ligusticum, etc.; angeliceae: Angelica,

Agasyllis, Chymsydia, etc.; peucedaneae: Myrrhi-

dendron, Diplotaenia, Johrenia, Cymbocarpum, Ferula,

Ferulago, Dorema, Opopanax, Peucedanum, Anethum,

etc.; tordylieae: Pastinaca, Heracleum, Tordylium,

Malabaila, Stenotaenia, Zosima, etc.: laserpitieae:

Laserpitium, Polylophium, Thapsia, Elaeoselinum, etc.

9.2 SANICULOIDEAE

Fruits with a soft parenchymatous endocarp, but contain-

ing scattered druses. Base of the stylodia surrounded by

a ring-like disc. – saniculeae: Sanicula, Polemannio-

psis, Steganotaenia, Hacquetia, Astrantia, Actinolema,

Alepidea, Eryngium; lagoecieae: Lagoecia, Petagnaea.

9.3 AZORELLOIDEAE

Leaves compound, stipulate. Fruits dorsally ﬂ attened,

with woody endocarp; n = 8. – Azorella, Bowlesia,

Diplaspis, Oschatzia, Homalocarpus, Drusa, Huanaca,

Spananthe, Diposis, Mulinum, Schizeilema, Laretia,

Dichosciadium, Dickinsia, Hermas, Choritaenia,

Bolax, Pozoa, Asteriscium, Gymnophyton, Eremocharis,

Domeykos, Stilbocarpa; Homalosciadium, Trachy-

mene, Uldinia, Klotzschia, Naufraga, Schoenolaena,

Pentapeltis, Chlaenosciadium, Brachyscias.

9.4 MACKINLAYOIDEAE

Woody or herbaceous plants. Axial parenchyma apo-

traceal or paratracheal. Leaves palmately compound

(Mackinlaya). Inﬂ orescences panicles or racemes,

often of umbels; carpophore present. Fruits laterally

ﬂ attened. – Apiopetalum, Mackinlaya, Centella,

Micropleura, Actinotus, Platysace, Xanthosia.

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