Takhtajan Armen. Flowering Plants

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Superoder CORNANAE 443

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Order 107. BRUNIALES

Shrubs or subshrubs, or rarely small trees, mostly with

ericoid habit, with subepidermal core. The hairs are

mostly long, slender and unicellular, with thick smooth

walls and narrow lumina. Resinous deposits observed

in cork, phloem, medullary rays and pith of Audoinia

and Lonchostoma. Vessels with scalariform perfora-

tions that usually have numerous bars. Fibers with

bordered pits. Axial parenchyma scanty paratracheal.

Rays heterogeneous, mixed uniseriate and pluriseriate.

Nodes unilacunar with one trace. Leaves evergreen,

alternate, simple, densely set, mostly subtrigonous

444 Subclass VII. ASTERIDAE

and acerose, often imbricate, entire, estipulate or

with small, rudimentary stipules, water-storage tissue

reported to occur at the apex of the leaf near the termi-

nations of the veins. Stomata anomocytic. Flowers in

dense spicate or capitate, terminal or axillary inﬂ ore-

scences, occasionally solitary, sometimes subtended by

several imbricate bracts, mostly small, bisexual, acti-

nomorphic. Sepals (4)5, free almost to the base, imbri-

cate, persistent. Petals (4)5, sessile or unguiculate, free

or connate into a variably long tube, imbricate, often

persistent. Stamens as many as petals and alternating

with them, often persistent; ﬁ laments free or sometimes

adnate to the corolla tube, in Lonchostoma anthers sub-

sessile and borne on the corolla tube, incurved in bud;

anthers small or very small, tetra sporangiate, dorsiﬁ xed,

often versatile, introrse, sometimes with a prolonged

connective, opening longitudinally. Intrastaminal nectary

disc sometimes present. Pollen 3-aperturate, or 6–11

aperturate, 3–10-colporate, tectate-columellate, almost

psilate to rugulose and murate-reticulate. Gynoecium

of two carpels, in Mniothamnea and Berzelia seemingly

unicarpellate or at least unilocular, in Audouinia with

three carpels, stylodia almost free to connate nearly to

the top into a style with stylar branches standing

close together; each stylodium or each stylar branch

with capitate stigma; ovary usually semi-inferior

or sometimes inferior, rarely almost superior, with

(1)2–4(-12) ovules per carpel. Dependent on the

superior or almost inferior position of the ovary,

nectarostomata are scattered on the ovary ﬂ anks, or

accumulated on the small hypgeneous part of the ovary

(Quint and Classen-Bockhoff 2006b). Ovules pendu-

lous from the summit of the septum or near the top

of the central axis of the ovary. Ovules anatropous,

epitropous, unitegmic, crassinucellate, with ventral

raphe. Female gametophyte of Polygonum-type. Fruit

dry, commonly crowned by the persistent calyx, often

indehiscent and achenelike or nutlike, 1-seeded, or the

1–2-seeded carpels separating and opening along

the ventral suture. Seeds with thin, exotestal seed

coat, sometimes arillate, with minute, straight well

differentiated embryo at the base of copious, ﬂ eshy

endosperm. Present proanthocyanidins and myricetin.

n = 10–11, 16 (Staavia), (21, 23).

Evidently belong to the Asteridae, probably related

to the Desfontainiales. According to Albach et al.

(2001) and Bremer et al. (2001, 2002), Bruniaceae are

sister to the Neotropical families Collumelliaceae and

Desfontainiaceae.

1. BRUNIACEAE

Berchtold et J. Presl 1820 (including Berzeliaceae

Nakai 1943). 12/75. Southern coastal regions of South

Africa, almost completely restricted to the Cape

Province.

Classiﬁ cation after M. Quint and R. Classen-

Bockhoff (2006a).

linconieae: Linconia; audouinieae: Audou inia,

Thamnea, Tittmannia; brunieae: Raspalia, Nebelia,

Staavia, Pseudobaeckea, Mnio thamnea, Brunia,

Lonchostoma, Berzelia.

Bibliography

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Bruniaceae. Bull. Torrey Bot. Club 117: 420–428.

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1–34.

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Linn. Soc. 96: 285–296.

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Lonchostoma and anderen Gattungen der Bruniaceae.

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121–206.

Quint M. 2004. Evolution of Bruniaceae: evidence from molec-

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of Mainz, Göttingen.

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Bruniaceae based on matK and its sequence data. Int. J. Plant

Sci. 167: 135–146.

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diversity and nectary uniformity in Bruniaceae. Bot. J. Linn.

Soc. 150: 459–477.

Superoder CORNANAE 445

Saxton WT. 1910. The ovule of the Bruniaceae. Trans. Roy. Soc.

South Afr. 2: 27–31.

Scott G. 1999. A chemosystematic and cladistic study of the

Southern African endemic family Bruniaceae. Ph.D. disser-

tation, University of Cape Town.

Weberling F. 1976. Weitere Untersuchungen zur Morphologic

des Unterblattes bei den Dikotylen: IX. Saxifragaceae s. 1.,

Brunelliaceae, und Bruniaceae. Beitr. Biol. Pﬂ . 52: 163–181.

Order 108. LOASALES (HYDRANGEALES)

Trees, shrubs, sometimes lianas, subshrubs, or peren-

nial or annual herbs. Indumentum consists of a wide

variety of characteristic trichomes. Woods range from

relatively soft to quite succulent. Vessels with scalari-

form, scalariform and simple, or (Loasaceae) basically

with simple perforations, but altered versions of sca-

lariform plates or plates with other aberrant conﬁ gura-

tions, lateral pitting various, sometimes laterally

elongate. Fibers with large or small bordered pits. Rays

usually heterogeneous to homogeneous, rarely almost

wanting. Axial parenchyma basically diffuse or vasi-

centric, rarely absent. Sieve-element plastids are of

S-type. Nodes mostly trilacunar. Leaves alternate or

opposite, simple, entire or variously divided, estipulate

or pseudostipulate. Stomata anomocytic or paracytic.

Flowers in cymose or rarely racemose inﬂ orescences,

spicate or capitate, less often solitary, usually bisexual,

actinomorphic. Sepals 4–5 (rarely 6–12), mostly con-

nate into a tube, imbricate or valvate, persistent. Petals

4–5 (rarely 6–12), free or sometimes more or less con-

nate, imbricate, contorted or valvate, Stamens (1)2 to

several, or numerous (up to 300), free or basally con-

nate into a short tube or fascicled or adnate to the

corolla tube, anthers basiﬁ xed or dorsiﬁ xed, versatile,

introrse or latrorse, tetrasporangiate, opening longitu-

dinally. Tapetum secretory. Microsporogenesis simul-

taneous. Pollen grains 2-celled or 3-celled, 3-colpate

or 3-colporate. Gynoecium of (2)3–5(-12) usually

united carpels; stylodia free or shortly connate below;

ovary mostly inferior or nearly inferior, plulocular to

1-locular, with one to several, or numerous, anatropous

to hemitropous, pendulous or ascending ovules in

each locule, or in each intruded parietal placenta.

Ovules unitegmic, tenuinucellate, rarely (Loasaceae

– Petalonychoideae and Gronovioideae) crassinucel-

late, with endothelium. Female gametophyte of

Polygonum-type. Endosperm cellular, with micropylar

and chalazal haustoria. Fruits septicidal or loculicidal

capsules, often ribbed, or sometimes indehiscent. Seeds

often minute, winged or not, exotestal, with straight or

curved embryo; endosperm copious, ﬂ eshy and oily,

or (Gronovioideae) endosperm scanty or wanting.

Producing various iridoid compounds (Weigend et al.

2000; Rodriguez et al. 2002), proanthocyanidins,

ﬂ avonols, and coffeic acids; n = 6–18 to 37.

The Loasales probably originated from the cunon-

ialean stock. However, they are only distantly related

to the Cunoniales (see Morgan and Soltis 1993).

Key to Families

1 Tanniniferous. Evergreen or deciduous, small trees

or shrubs, sometimes lianas, subshrubs, or rhizoma-

tous herbs. Indumentum of stellate hairs, long, uni-

cellular trichomes or (Broussaisia) tufted trichomes.

Often with raphid sacs in both leaf and axis. Vessels

typically with scalariform perforations that have

20–90 thin bars, but sometimes with both scalari-

form and simple plates or rarely even all simple;

lateral pitting scalariform to opposite or rarely alter-

nate. Fibers typically with large or small bordered

pits, but sometimes with simple pits, sometimes

septate. Rays heterogeneous or less often homoge-

neous. Axial parenchyma absent, or more often dif-

fuse and scanty vasicentric. Nodes trilacunar or

sometimes multilacunar. Leaves generally opposite,

entire or dentate, or sometimes lobed, glandular

leaf teeth occur in Decumaria, Deutzia, and

Philadelphus, estipulate. Stomata anomocytic or

paracytic. Inﬂ orescences cymose or corymbose,

sometimes heads, or racemes by abortation, rarely

ﬂ owers solitary. Flowers usually conspicuous, all

equal or marginal ones sterile and with enlarged

sepals, bisexual or rarely (Broussaisia) polygamo-

dioecious. Sepals 4–5(-12), mostly connate into a

tube, lobes valvate or imbricate. Petals 4–5(-12),

free, valvate, imbricate or contorted. Stamens

(1)2-several times as many as petals, sometimes as

many as 50 or even (Carpenteria) up to 200; ﬁ la-

ments free or slightly connate at base; anthers bas-

iﬁ xed or dorsiﬁ xed, versatile, introrse or latrorse.

Pollen grains 3-colpate (some species of Hydrangea)

or 3-colporate. Gynoecium of (2)3–5(-12) carpels;

stylodia free or shortly connate below, the branches

often stigmatic for much of their length or seldom

the style short, with distinct stigmas or a capitate,

nearly lobed stigma; ovary inferior or less often

446 Subclass VII. ASTERIDAE

superior or semi-inferior, plurilocular to unilocular,

with one to several or numerous, anatropous,

pendulous or ascending ovules in each locule or on

each intruded parietal placenta. Epigynous disk

usually present. Fruits septicidal or more often loc-

ulicidal capsules or sometimes (Dichroa,

Broussaisia) berries. Seeds winged or not; embryo

large, straight, surrounded by ﬂ eshy endosperm,

sometimes with aril (Whipplea, Fendlerella –

Hufford 1995; Lobova 2000); exotestal cell signi-

ﬁ ed. Present rare iridoid compounds (deutziosideas),

seco-iridoids and loganin, proanthocyanidins (cyani-

din or delphinidin), ﬂ avonols (kaempferol, myrice-

tin, ﬂ avonols), n = 13–18.. . . . .1. hydrangeaceae.

1 Non-tanniniferous perennial or annual herbs, coarse,

erect or twining, less often shrubs or very rarely

(Mentzelia arborescens) small and succulent trees

or (Fuertesia) woody lianas. Indumentum consists

of a wide variety of characteristic trichomes, some

of which are elongated, siliciﬁ ed, bristly stinging

hairs ﬁ lled with a yellowish irritant substance in the

hairs themselves or the surrounding basal cells; the

irritant substance is readily liberated owing to the

brittleness of the hairs; glandular hairs with uniseri-

ate stalks and uniseriate heads frequently contain

cystoliths. Woods range from relatively soft to quite

succulent. Vessels basically with simple perfora-

tions, but sometimes (in Loasa picta and Mentzelia

humilis) altered versions of scalariform plates or

plates with other aberrant conﬁ gurations were

observed by Carlquist (1984); lateral pitting vari-

ous, sometimes laterally elongate or even scalari-

form-like. Fibers with bordered pits, but sometimes

with vestigial borders on pits. Rays tall and wide,

predominantly multiseriate, heterogeneous to

homogeneous, rarely almost wanting. Axial paren-

chyma basically diffuse, rarely absent

(Presliophytum), or vasicentric (Plakothira frute-

scens). Leaves alternate or opposite, simple, entire

or variously divided, estipulate or pseudostipulate.

Stomata anomocytic. Flowers in cymose or rarely

racemose inﬂ orescences, spicate or capitate, less

often solitary, bisexual, actinomorphic. Sepals 4–5

(rarely 6–7), connate; calyx tube adnate to the ovary,

often ribbed, the ribs sometimes spirally twisted;

lobes imbricate or valvate, persistent. Petals 4–5

(rarely 6–7), inserted on the calyx, sessile or clawed,

imbricate, contorted or valvate, free or sometimes

more or less connate. Stamens commonly numer-

ous (up to 300), less often 10 in two cycles or only

ﬁ ve or only two fertile, free or basally connate

into a short tube or fascicled or adnate to the

corolla tube, sometimes with nearly sessile anthers

on the corolla tube; some of the stamens often mod-

iﬁ ed into petaloid, scalelike, or nectariferous sta-

minodia; anthers basiﬁ xed. Pollen grains

3-colporate, tectate-columellate, with striate-reticu-

late, striate, or, more rarely, shallowly reticulate

tectum. Gynoecium of 5–3, rarely of 6–7 united

carpels, with simple style; ovary inferior or nearly

inferior, 1-locular often with more or less deeply

intruded parietal placentas, rarely fully partitioned,

with 1-many ovules on each placenta; in

Gronovioideae pseudomonomerous, with a solitary

pendulous apical ovule. Ovules anatropous to

hemitropous, unitegmic with a fairly long micro-

pyle, tenuinucellate, with endothelium; in

Gronovioideae and Petalonychoideae ovules crassi-

nucellate, in many genera ovules with cup-shaped

funicula-projections. Female gametophyte of

Polygonum-type. Endosperm cellular, with aggres-

sive micropylar and chalazal haustoria. Fruits septi-

cidal or loculicidal capsules, often ribbed, or

sometimes indehiscent. Seeds often minute, exot-

estal, with straight or curved embryo surrounded by

copious, oily, thin-walled endosperm or

(Gronovioideae) endosperm scanty or wanting.

Producing quercetin, coffeic acid, and verious iri-

doid compounds (Weigend et al. 2000; Rodriguez

et al. 2002), n = 6–15 to 37. . . . . . . 2. loasaceae.

1. HYDRANGEACEAE

Dumortier 1829 (including Hortensiaceae Berchtold

et J. Presl 1820; Kirengeshomaceae Nakai 1943;

Philadelphaceae Martynov 1820). 17/250. Subtropical

and warm-temperate regions of the Northern Hemi-

sphere, especially eastern Asia (mainly China) and

North America; a few species in Southeast Asia and in

the Andes, from Mexico to southern Chile.

1.1 PHILADELPHOIDEAE

Indumentum mostly of stellate hairs. Flowers in termi-

nal cymes or racemes, rarely solitary. Filaments often

with elongate lateral teeth at apex. – philadelpheae:

Carpenteria, Fendlerella, Fendlera, Jamesia, Philade-

lphus, Whipplea; deutzieae: Deutzia.

Superoder CORNANAE 447

1.2 HYDRANGEOIDEAE

Indumentum of simple hairs or absent. Flowers in

cymose inﬂ orescences, sometimes corymbose or

capitate, or pseudoracemose by reduction. Filaments

not toothed. – hydrangeae: Hydrangea, Platycrater,

Decumaria, Pileostegia, Schizophragma; DICHROEAE:

Dichroa, Broussaisia; cardiandreae: Cardiandra,

Deinanthe.

1.3 KIRENGESHOMOIDEAE

Indumentum of 2-armed hairs. Leaves palmatiﬁ d. Flowers

in tornately branched terminal and axillary paniculate

cymes. Filaments not toothed. – Kirengeshoma.

The monotypic genus Kirengeshoma (southern Japan,

Korea, temperate China) occupies a rather isolated

position. Typically, it strongly differs from other mem-

bers of the family in that its indumentum consists of

2-armed hairs; it has 15 stamens in three cycles, its three

antipodal cells become binucleate and function as haus-

toria, and its seeds are winged and anatomically differ

from those of the rest of Hydrangeaceae (Krach 1976).

Besides, its pollen grain has a unique complete tectum

associated with partial tectum (Hideux and Ferguson

1976). However, according to Hideux and Ferguson, a

transitional state between a complete tectum and perfo-

rate tectum (a sparsely, perforated, unsculptured tectum)

occurs in Anopterus macleyanus and Polyosma. Also,

Kirengeshoma has many features in common with the

tribe Cardiandreae, and serologically it is very close to

Hydrangea (Grund and Jensen 1981).

2. LOASACEAE

A.L. de Jussieu 1804 (including Cevalliaceae Grise-

bach 1854; Gronoviaceae Endlicher 1841). 20/300.

Temperate, subtropical, and tropical regions of America

from southwestern Canada to Argentina and Chile, and

West Indies, Galapagos Islands, with only two genera

in the Old World – Plakothira (Marquesas) and Fissenia

(F. capensis in southwestern Africa and F. arabica in

Somalia, northeastern Ethiopia, and southern and south-

western parts of the Arabian peninsula).

Classiﬁ cation after M. Weigend (2004).

2.1 LOASOIDEAE

Leaves opposite below and alternate above or opposite

throughout; stinging hairs sometimes present. Fertile

stamens numerous, arising with centrifugal matura-

tion, rarely fewer and opposite the petals. Staminodia

absent. Ovules numerous to three, on parietal

(sometimes subapical) placentas, n = 6. – loaseae:

Chichicaste, Huidobria, Presliophytum, Aosa,

Blumenbachia, Loasa, Caiophora, Scyphanthus, Nasa;

kissenieae: Fissenia (Kissenia); klaprothieae:

Klaprothia, Plakothira, Xylopodia.

2.2 MENTZELIOIDEAE

Leaves opposite below and alternate above, with

scabrid-glochidiate trichomes, sometimes with sting-

ing hairs. Stamens numerous, arising with centripetal

maturation. Staminodia, if present, petaloid and free to

base. Ovules three to numerous, n = 7. – Mentzelia,

Eucnide, Schismocarpus.

2.3 GRONOVIOIDEAE

Leaves always alternate, always with glochidiate

trichomes, sometimes with stinging hairs. Inﬂ orescences

terminal thyrsoids. Fertile stamens 4–5, alternating

with the petals. Staminodia absent. Ovules solitary,

pendulous from an apical placenta. – Cevallia,

Gronovia, Fuertesia.

2.4 PETALONYCHOIDEAE

Leaves alternate with glochidiate hairs with short, spread-

ing branches, without stinging hairs. Inﬂ orescences termi-

nal racemes. Fertile stamens reduced to two. – Petalonyx.

Related to the Hydrangeaceae, see Fan and Xiang

(2003) and Weigend (2004), Soltis et al. (2006).

Bibliography

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Order 109. CORNALES

Trees or more often shrubs, sometimes ericoid shrubs

(Grubbiaceae), rarely rhizomatous herbs. Hairs simple,

unicellular. Vessels mostly with scalariform perfora-

tions usually with many bars; lateral pitting scalari-

form or opposite. Fibers typically with large, often

conspicuous bordered pits. Rays markedly heteroge-

neous. Axial parenchyma typically apotracheal. Sieve-

element plastids of Ss-, or less often So-, and Pcs-types.

Nodes trilacunar. Leaves alternate or more often oppo-

site, simple, mostly entire, estipulate. Stomata usually

anomocytic. Flowers small, in various types of cymose

inﬂ orescences, usually bisexual, actinomorphic or very

rarely slightly zygomorphic, 4-merous or less often

5-merous. Calyx adnate to ovary, 4–10 lobes or teeth

or obsolete. Petals free or rarely basally connate, imbri-

cate or valvate, reduced or wanting in female ﬂ owers.

Stamens as many as and alternate with petals or less

often twice or even three or four times as many as the

petals, mostly attached to or around the edge of epigy-

nous nectary disc; ﬁ laments free; anthers tetrasporan-

giate, opening longitudinally. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains

2-celled, 3-colporate or of 3-colporate-derived type.

Nectary disc usually present. Gynoecium of 2–9

(mostly two) united carpels; stylodia more or less free

or united into a style with a lobed or capitate stigma;

ovary inferior, with one pendulous ovule per locule.

Ovules anatropous or rarely (some species of Cornus)

hemitropous, apotropous, or epitropous, unitegmic,

crassinucellate or (Nyssaceae and some species of

Cornus) tenuinucellate, often with endothelium.

Female gametophyte usually of Polygonum-type,

rarely (some species of Cornus) tetrasporic and then

usually of Fritillaria-type. Endosperm cellular

(Davidiaceae, Nyssa aquatica, Alangium chinense,

Cornaceae, Grubbiaceae) or nuclear (Nyssaceae,

Alangium lamarckii). Fruits drupes. Seeds small to

medium-sized, exotestal, with small to rather large

embryo; endosperm usually copious. Usually contain

iridoid compounds, mostly cornin, and gallic and

ellagic acids (and corresponding tannins).

Cornales are so close to the Loasales, that the

boundary between them is almost conventional. The

most archaic cornalean family is Davidiaceae.

Key to Families

1 Gynoecium of (6)7(-10) carpels. Nectary disc

absent. Deciduous trees. Vessel elements very long,

with very slanting end walls and scalariform perfo-

rations with 60–100 thin bars; lateral pitting oppo-

site or less often transitional. Fibers with distinct

bordered pits in both radial and tangential walls.

Axial parenchyma diffuse(-in-aggregates). Rays

heterogeneous with long ends. Crystals absent.

Leaves alternate, cordate at base, acuminate, ser-

rate-dentate, pinnately veined with 3–7 subbasal

veins, usually silky-pubescent below when young.

Flowers andromonoecious, apetalous, in terminal

globose heads subtended by two (rarely three) large,

creamy-white, subopposite spreading bracts; male

ﬂ owers numerous, scarcely distinct in dense head,

bisexual ﬂ ower solitary in the head, laterally placed

450 Subclass VII. ASTERIDAE

obliquely above the middle of the axis. Male ﬂ ow-

ers completely devoid of perianth and consist of

(1-)5–6(-12) stamens with slender, elongate ﬁ la-

ments and small introrse anthers. Pollen grains

3-colporate. Bisexual ﬂ owers: calyx adnate to the

ovary and with much reduced limb; stamens 15–26,

mostly 21, arranged around the base of style; style

short, thick, conical, with 6–10 spreading stigmas;

stigmas sulcate and papillose on their adaxial side;

ovary 6–10-locular, with one ovule per carpel.

Endothelium present. Fruits drupaceous, with

granular mesocarp and bony longitudinally sulcate

(1–2) 3–5-seeded endocarp. Seeds with large

embryo subequal to fleshy, oily endosperm.

Contain gallic and ellagic acids (and correspond-

ing tannins), n = 20, 21. . . . . . . . 1. davidiaceae.

1 Gynoecium of two or less often (Curtisiaceae) four

carpels. Nectary disc present. Petals mostly present.

2 Petals (when present) imbricate, small. Trees and

shrubs. Hairs simple, unicellular, sometimes with

granular surface, together with variously shaped

unicellular glands on leaves. Vessels with sca-

lariform or scalariform-reticulate perforations

that have numerous (17–50) thin bars; lateral pit-

ting opposite or scalariform. Fibers with dis-

tinctly bordered pits in both radial and tangential

walls, occasionally (in some species of Nyssa)

septate, or with dark purple deposits. Rays het-

erogeneous, with short ends. Axial parenchyma

diffuse, but in Nyssa occasionally scanty paratra-

cheal. Prismatic crystals present (Noshiro et Baas

1998). Leaves alternate, entire or denticulate,

pinnately veined. Flowers small, greenish-white,

polygamous, polygamo-dioecious, or dioecious,

in terminal panicles of globose heads

(Camptotheca) or (Nyssa) male ﬂ owers crowded

in axillary, short racemes, umbels, or heads,

bisexual and female ﬂ owers in few-ﬂ owered

heads or solitary by reduction. Calyx represented

by a minute, 5-dentate or irregularly toothed rim,

or virtually obsolete. Petals ﬁ ve or rarely more,

small, imbricate, in female ﬂ owers sometimes

wanting. Stamens in male ﬂ owers (5-) 10(16),

often 2-seriate; ﬁ laments elongate, subulate;

anthers small, introrse or latrorse. Pollen grains

3-colporate. Nectary disc in male ﬂ owers ﬂ eshy,

pulvinate. Rudimentary gynoecium wanting, or

less often there is a very small subulate rudiment

in the middle of the disc. Stamens in bisexual

ﬂ owers as many and alternate with the petals or

wanting; anthers fertile or sterile. Gynoecium

usually pseudomonomerous; 1-locular, with one

pendulous ovule, but in Nyssa there sometimes

occurs a gynoecium with a 2-locular ovary and

two free stylodia; ovary is crowned at the top

with broad pulvinate or conical nectary disc in

the middle of which stands a recurved stylodium

of the fertile carpel with decurrent stigma on its

inner side (Nyssa) or a short, 2-ﬁ d style

(Camptotheca). Endothelium absent. Fruits

drupaceous (Nyssa) or shortly subsamaroid

(Camptotheca). Seeds with short, straight embryo

and copious endosperm; radicle short. Contain

gallic and ellagic acids, alkaloid camptothecin

(Camptotheca), and corresponding tannins, n =

21 (Camptotheca), 22 (Nyssa). . . 2. nyssaceae.

2 Petals valvate.

3 Leaves strongly serrate, more or less ﬂ at.

Evergreen trees. Branchlets, petioles, and inﬂ o-

rescences with stellate hairs. Resin ducts absent.

Wood diffuse-porous. Vessels with scalariform

perforations that have 26–39 bars; lateral pit-

ting opposite, without vestures. Fibers with dis-

tinctly bordered pits. Rays heterogeneous.

Axial parenchyma diffuse to diffuse-in-aggre-

gates. Prismatic crystals common in ray cells

(Noshiro et Baas 1998). Leaves opposite,

coarsely dentate, coriaceous, shiny above, more

or less ferrugineo-tomentose below. Flowers

minute, in many-ﬂ owered, terminal trichoto-

mously branched tomentose thyrses, bisexual

or rarely (Afrocrania) dioecious, bracteolate.

Calyx tube turbinate, 4-lobed, lobes open.

Petals four, triangular-ovate. Stamens four,

alternipetalous; ﬁ laments subulate; anthers cor-

date at the base, introrse. Pollen grains small,

3-colporate, tectum complete with pseudoper-

forations, or H-shaped. Intrastaminal disc

broad, 4-angled, densely barbate. Ovary infe-

rior, 4-locular, ovule one per locule. Style short,

subconical, glabrous, with 4-lobed stigma;

Fruits small, subglobose, 4-locular, 4-seeded

drupes arcolate at the apex, crowned by the

calyx. Seeds elongate- oblong, subcylindric,

with elongate embryo in copious endosperm.

Contain tannins, proanthocyanins, ellagitannins

and iridoids, n = 13.. . . . . . . .4. curtisiaceae.

3 Leaves entire.

4 Inﬂ orescences capitate or cone-like, compound

of dichasia. Evergreen ericoid shrubs; hairs

Superoder CORNANAE 451

unicellular. Leaves opposite-decussate, estipu-

late. Flowers in small axillary 3(2)-ﬂ owered

dichasia or in many-ﬂ owered conelike com-

pound dichasia, the ovaries coherent or connate.

Flowers subtended by bracts and provided with

two prophylls. Sepals four, free, valvate, hairy

on outer surface. Petals absent. Stamens eight,

half of them alternate with the sepals, and the

other longer ones are opposite and slightly

adnate to their bases; anthers mediﬁ xed or bas-

iﬁ xed, inverted, opening longitudinally along

the lateral C-like line; peripheral microsporan-

gia nonfunctional, vestigial. Pollen grains

3-colporate, psilate. Disc epigynous, papillate

to shortly hairy. Gynoecium of two carpels;

style unbranched, with two short apical lobes;

ovary inferior, at ﬁ rst 2-locular, but later

becoming incompletely 1-locular, with two

ovules pendulous from the summit of the sep-

tum. At the cotyledonary stage the massive cel-

lular endosperm shows chalazal and micropylar

haustoria (Fagerlind 1947). Fruits 1-seeded

drupes that connate forming a dense frutes-

cence. Seeds with thin seed coat and large,

straight, cylindrical embryo surrounded by the

copious, oily endosperm. Contain tannins but

there are negative reports of ﬂ avonoids and iri-

doids (Kubitzki 2004). . . . . . 5. grubbiaceae.

4 Inﬂ orescences thyrso-paniculate.

5 Ovules epitropous (with a ventral raphe and

micropyle directed upwards). Evergreen trees.

Resin ducts with secretory epithelium present

in vegetative organs and in fruits. Hairs uni-

cellular, two-armed. Wood diffuse-porous.

Vessels with scalariform perforations that have

numerous bars (34–66); lateral pitting oppo-

site-scalariform, without vestures. Fibers with

distinctly bordered pits in both radial and tan-

gential walls. Rays heterogeneous with long

ends. Axial parenchyma apotracheal in narrow

bands. Crystals absent (Diplopanax) or

(Mastixia) prismatic crystals occasionally

present in body ray cells (Noshiro et Baas

1998). Leaves alternate or opposite, entire,

coriaceous, pinnately veined. Flowers small,

usually in terminal many-ﬂ owered compound

dichasia, bisexual; pedicels 2-bracteolate,

articulated. Calyx campanulate, 4–5-dentate,

open. Petals 4–5, coriaceous, inﬂ exed at the

apex and ﬁ mbriate or 2-dentate. Stamens 4–5,

alternipetalous, inserted at the base of ﬂ eshy,

intrastaminal tetragonous or pentagonous

nectary disc; ﬁ laments short; anthers introrse.

Pollen grains 3-colporate, sometimes 4-colpo-

rate, with complete or partially perforate tec-

tum. Gynoecium pseudomonomerous; style

short, thick, with punctiform or rarely 2-lobed

stigma; ovary 1-locular. Fruit drupes with

grooved endocarp. Seeds with small embryo in

copious endosperm. Contain gallic and ellagic

acids (and corresponding tannins), unidentiﬁ ed

iridoids reported from leaf tissue (Bate-Smith

et al. 1975), n = 11, 13. . . . . 3. mastixiaceae.

5 Ovules apotropous (with a dorsal raphe and

micropyle directed downward).

6 Leaves opposite or rarely (in Swida) alternate.

Evergreen or deciduous trees and shrubs, rarely

rhizomatous herbs with annual stems, provided

with characteristic two-armed hairs. Plants

without laticifers. Vessels with scalariform

perforations that usually have 14–71 bars.

Fibers with distinctly bordered pits in both

radial and tangential walls, without vestures.

Rays heterogeneous. Axial parenchyma usu-

ally diffuse or diffuse-in-aggregates. Crystals

rare: prismatic crystals in Cornus peruviana,

together with druses in some species (Noshiro

et Baas 1998). Leaves entire. Flowers small, in

terminal, cymose inﬂ orescences, paniculate,

corymbose umbellate or capitate (the latter

sometimes subtended by a whorl of large peta-

loid bracts), bisexual or rarely (Afrocrania)

dioecious. Calyx with four small teeth around

the summit of the ovary or entire. Petals four.

Stamens four, alternipetalous, around the edge

of pulvinate, intrastaminal nectary disc; anthers

oblong or ellipsoid, dorsiﬁ xed. Pollen grains

3-colporate, with H-shaped columellae or tec-

tum complete with short supratectal spines, of

which at least some are prolongations of the

columellae, sometimes perforate. Style ﬁ liform

to columnar, with capitate or truncate stigma;

ovary 2-locular, with one ovule per locule.

Fruits drupes, areolate at apex, with bony or

crustaceous, 2-locular endocarp. Seeds oblong,

compressed, with membranous testa, elongate

embryo and ﬂ eshy endosperm; seed-coat thin-

walled and crushed. With iridoid compounds

but without isoquinoline alkaloids and petrose-

linic acid. Contain gallic and ellagic acids (and

452 Subclass VII. ASTERIDAE

corresponding tannins), n = 9–11, or (C.

canadensis) 22.. . . . . . . . . . . . .6. cornaceae.

6 Leaves alternate. Small trees or less often

shrubs or woody lianas, sometimes thorny.

Hairs somewhat diverse, sometimes stellate and

glandular; the Madag ascan species Alangium

grisolleoides has 2-armed hairs. Articulate

laticifers present in various tissues of the shoot.

Vessels mostly with simple perforations, but in

section Constigma they are scalariform with

7–16 bars; lateral pitting alternate and dense.

Fibers mostly with minutely bordered pits, or

with simple pits and slit-like abertures in sec-

tion Constigma. Rays heterogeneous with long

ends. Axial parenchyma diffuse or diffuse-in-

aggregates. Prismatic crystals usually conspic-

uous in both ray cells and axial parenchyma

(Noshiro et Baas 1998). Leaves entire or lobed,

pinnately or palmately veined, often some of

the cells of the mesophyll containing cluster

crystals and appearing as translucent dots.

Flowers in axillary cymes, bisexual or rarely

unisexual (A.grisolleoides dioecious). Calyx

with 4–10 lobes or teeth or truncate. Petals

4–10, linear or strap-shaped, valvate or rarely

contorted, hairy within, sometimes connate at

base, ﬁ nally recurved or revolute. Stamens

4–40, arranged in a single cycle around a con-

spicious, pulvinate, or sometimes almost hemi-

spherical nectary disc; ﬁ laments free or slightly

connate at base, sometimes adnate to the petals

at base, more or less barbate or villose on inner

side; anthers more or less linear, introrse, bas-

iﬁ xed or more rarely dorsiﬁ xed and versatile

and widely sagittate. Pollen grains 3–4(-8)-col-

porate or porate. Style elongate, ﬁ liform with

clavate, 2–4-lobed or more or less capitate

stigma or rarely style short and with two elon-

gate, ventrally stigmatic branches; ovary 2-loc-

ular or more often 1-locular. Endothelium

present. Fruits drupes crowned by calyx and

disc and with crustaceous or woody endocarp,

1-locular or less often 2-locular, each locule

with one seed or one locule empty. Seeds with

large, straight embryo, foliaceous cotyledons,

and elongate hypocotyl; endosperm copious,

oily, and also with reserves of hemicellulose;

seed-coat thin-walled. Without iridoid com-

pounds but commonly producing isoquino-

line (emetine) alkaloids (alangiside and

tubulosin), n = 8, 9, 11.. . . .7. alangiaceae.

1. DAVIDIACEAE

H.L. Li 1955. 1/1. Southwestern and central China.

Davidia

In spite of very specialized inflorescences and

reduced perianth, Davidia is characterized by very

primitive wood structure and multicarpellate gynoe-

cium. We may therefore agree with Eyde (1967:

167) that «with respect to these characters Davidia

is the nearest living thing to the common ancestor

of Cornus and Nyssa». Usually Davidia is included

in Nyssaceae, but from Nyssa and Camptotheca it

differs not only in morphology of inflorescences

and flowers (including the absence of the nectary

disc), crassinucellate ovules with endothelium and

some other embryological characters (Kamelina

and Shevchenko 1988) but also palyno logically

(Eramian 1971), chemically (Bate-Smith et al.

1975), and serologically (Fairbrothers 1977).

2. NYSSACEAE

A.L. de Jussieu ex Dumortier 1829. 2/9. Eastern

Himalayas, Assam (Khasi Hills), continental China,

Southeast Asia, eastern North America. Camptotheca

(1) is endemic to eastern Tibet and southwestern and

central China.

Nyssa, Camptotheca

Related to the Davidiaceae but in some respects

more advanced.

3. MASTIXIACEAE

Calestani 1905. 2/c.20. Sri Lanka, southwestern and

northeastern India, eastern Himalayas, northern Burma,

southwestern and southern China, Hainan Indochina,

Andaman Islands, Malesia, New Guinea New Britain,

Solomon Islands. Diplopanax (2) is distributed in China

(Guangdong, Guangxi, Guizhou and Yunnan) and

northern Vietnam.

Diplopanax, Mastixia

Differ from the Nyssaceae and Cornaceae and

related families in chemical features (Bate-Smith et al.