Takhtajan Armen. Flowering Plants

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Superorder RANUNCULANAE 91

Troll W. 1933. Beiträge zur Morphologic des Gynoe- ceums: III.

Über das Gynoeceum von Nigella und einiger anderer

Helleboreen. Planta 21: 266–291.

Tucker SC. 1966. The gynoecial vascular supply in Caltha.

Phytomorphology 16: 339–342.

Tucker SC and SA Hodges. 2005. Floral ontogeny of Aquilegia,

Semiaquilegia and Enemion (Ranunculaceae). Int. J. Plant

Sci. 166: 557–574.

Vishnu-Mittre and BD Sharma. 1963. Studies of Indian pollen

grains 2. Ranunculaceae. Pollen et spores 5: 285–296.

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Asteropyrum (Ranunculaceae) inferred from chloroplast and

nuclear sequences. Plant Syst. Evol. 255: 41–54.

Wang XQ, DY Hong, and ZY Li. 1993. A study on pollen and

seed coat in the tribe Cimicifugeae and some allied genera

(Ranunculaceae). Cathaya 5: 131–149.

Wang XQ, ZR Deng, and DY Hong. 1998. The systematic

position of Beesia: evidence from ITS (nrDNA) sequence

analysis. Acta Phytotax. Sinica 36: 403–410.

Wang XQ, ZY Li, and DY Hong. 1994. A karyomorphological

studie of nine species in four genera of Ranunculaceae.

Cathaya 6: 43–56.

Werth E. 1941. Die Blütennektarien der Ranunculaceen und ihre

phylogenetische Bedeutung. Ber. Deutsch. Bot. Ges. 59:

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Eranthis and seven other genera of Ranunculaceae: evidence

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267–289.

Yuan Q and Q-E Yang. 2006b. Cytology, palynology and

taxonomy of Asteropyrum and four other genera of

Ranunculaceae. Bot. J. Linn. Soc. 152: 15–26.

Zhang ZY. 1982. Chromosome observation of three

Ranunculaceous genera in relation to their systematic posi-

tion. Acta Phytotax. Sinica 20: 402–409.

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family Ranunculaceae. Ann. Missouri Bot. Gard. 76:

1012–1049.

Order 26. CIRCAEASTERALES

Small perennial or annual herbs. Vessels usually with

simple perforations. Nodes unilacunar. Leaves alter-

nate or opposite, palmate or simple, 2-ranked, petio-

late, with an open dichotomous venation, the veins

ending in teeth of the lamina, estipulate. Stomata ano-

mocytic. Flowers very small, in terminal cyme

(Circaeaster) or solitary, devoid of bracts and bracte-

oles, bisexual, actinomorphic, apetalous. Sepals 2–3 or

5–9, scalelike or petaloid, valvate, persistent. Floral

nectaries present (Kingdonia) or absent (Circaeaster).

Stamens 1(2)3 or 14–17; anthers tetrasporangiate or

disporangiate, introrse or latrorse, opening longitudi-

nally; connective not prolonged. Tapetum glandular.

Pollen grains 2-celled, 3-colpate, with striate tectum.

Gynoecium apocarpous, of (1)2-9 conduplicate car-

pels with 1 or 2 pendulous ovules per carpel. Ovule

orthotropous (hemitropous according to Hu et al.

1990), unitegmic, tenuinucellate. Before fertilization

integument covers 1/3 part of nucellus, after that, elon-

gates and then disappers (Ren Y. et al. 1998). Female

gametophyte tetrasporic, 4- or 8-nucleate. Fertilization

porogamous (Kingdonia) or mesogamous (Cicraeaster)

Endosperm helobial or cellular. The ﬁ rst division

of primary endosperm nucleus longitudinal. The

embryogeny of Kingdonia subtype of Caryophyllad-

type. Fruits achenes. Seeds small, without testa, seed

coat degenerating, embryo relatively large. Endosperm

differentiated into two parts – storage micropylar part

and absorbing chalazal part, which are separated by an

intermediate suberized layer; the cells of endosperm

surface with thikend outer. n = 9, 15.

A rather isolated group that shows some similarities

with Lardizabalales, Berberidales and Ranunculales.

They often included in Ranunculales, but markedly

differ from them, testaless seeds and some embryo-

logical features such as: Caryophyllad type of

embryogeny, the ﬁ rst division of primary endosperm

92 Subclass II. RANUNCULIDAE

nucleus, differentiating of endosperm on two parts,

thickened outer walls of cells of endosperm surface,

integument covering 1/3 part of nucellus before fertil-

ization and disappearing after that (Ren Y. et al. 1998).

However, according to Nowicke and Skvarla (1982),

palynologically Circaeaster and Kingdonia are related

to Hydrastis and to the tribe Epimedieae in the

Podophyllaceae than to any remaining taxa examined in

the Ranunculanae. They base the relationships on the

similarity of the tectum, a compound layer of striae.

However, there are also many dissimilarities that make

Circaeasterales a very distinct group (Takhtajan 1997).

Key to Families

1 Perennial herbs with long, creeping, slender, branch-

ing, scaly rhizome with endotrophic mycorrhizae, a

solitary leaf and ﬂ ower arising from each bud.

Unilacunar node with four traces in foliage leaves and

a double trace in both cataphylls and scale leaves.

Leaves usually single, basal, long-petiolate, with

membranous, suborbicular lamina, 5–7-palmatisect,

segments cuneate, variously lobed and dentate on the

upper margin. Flowers solitary, with a long pedicel.

Sepals 5–6, rarely 8–9 in two series with 5–6 in the

outer and 3 in the inner series, pale yellow-green,

ovate, acuminate, with dichotomous venation.

Androecium consists of 9–11 outer sterile and 5–6

inner fertile stamens; each sterile stamen differenti-

ates into a stalk and a broadened head with a groove

in the centre; fertile stamens differentiate into costate

ﬁ laments and 2-locular, tetrasporangiate, extrorse

anthers (Ren et al. 2005). Pollen grains and the tec-

tum consisting of two layers of striae or mostly stri-

ate. Gynoecium of 5–6, rarely 9 carpels, opened in the

upper third, with subulate stylodium, which gradually

becomes recurved over the dorsal edge of the carpel,

and 1 ventral pendulous, hemitropous ovule per car-

pel. Inner epidermal cells of integument near the

micropyle elongated radically to form a palisade-like

structure. Endosperm helobial. Achenes 1–5, gla-

brous, with thick stipe, beaks elongate, strongly bent

outward at base. Seeds narrow, ellipsoid; endosperm

cell walls thin, n = 9. . . . . . . . . . . 1. kingdonia ceae

1 Annual herbs with an elongated stemlike hypocotyl,

short stem, and linear persistent cotyledons and leaf-

rosette. Unilacunar nodes with a single trace in foli-

age leaf. Leaves more or less opposite, petiolate,

cuneate-spatulate, dentate at apex. Inﬂ orescences are

thyrsoid with extremely short main axis and 10-30

(-40) ﬂ owers; 2–3 ﬂ owers in the center of the inﬂ o-

rescence have longer pedicels and bloom ﬁ rst (Tian

et al. 2006). Flowers bisexual, nearly actinomorphic.

Sepals 2–3, scalelike, valvate, persistent, without vas-

cular bundles. Stamens (1-)2(-3), alternisepalous;

anthers basiﬁ xed, disporangiate, latrorse. Pollen grains

with the irregularly striate tectum. Gynoecium of

1-2(-4) fusiform, shortly stipitate, closed carpels with

very short stylodium and slightly elongate and slightly

oblique papillate stigma. Ovule solitary, small to very

small, orthotropous, pendulous, submarginal, the

uppermost ovule aborting. Integument without pali-

sade-like structure. Endosperm cellular. Fruits long

elliptic green achenes covered by uncinate hairs. Seeds

small; embryo well differentiated, small, straight, with

short cotyledons; endosperm copious; endosperm cell

wall thick. n = 15. . . . . . . . . . . 2. circaeasteraceae

1. KINGDONIACEAE

A. S. Foster ex Airy Shaw 1964.1/1. High mountains

of southwestern China, primarily in western Sichuan.

Kingdonia.

2. CIRCAEASTERACEAE

Hutchinson 1926. 1/1. Northwestern Himalayas to

western China.

Circaeaster.

Despite some common features with the

Kingdoniaceae, such as dichotomous leaf venation,

pollen morphology, pendulous, unitegmic, tenuinucel-

late ovules. Circaeasteraceae have many unique char-

acters that make them rather distinct.

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Superorder RANUNCULANAE 93

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signiﬁ cance. Acta Phytotax. Sinica 20: 402–409.

Order 27. PAPAVERALES

Perennial or annual herbs or rarely shrubs

(Dendromecon and species of Bocconia) or small soft-

wooded trees up to 6 m tall. Latex present throughout

the plant in either laticiferous tubes or laticiferous

sacks or plants without latex and laticifers, but

with elongate idioblasts containing juice (except

94 Subclass II. RANUNCULIDAE

Pteridophyllum, which lacks both idioblasts and latic-

ifers). The transverse section of the stem mostly

exhibits a single ring of widely spaced collateral

vascular bundles, which are separated by broad multi-

seriate medullary rays (markedly heterogeneous in

Bocconia). Vessels with simple perforations. Fibers

with numerous, small, simple pits. Axial parenchyma

paratracheal. Nodes unilacunar to multilacunar. Leaves

alternate (sometimes all basal) or sometimes opposite

or almost verticillate, entire or more often variously

divided, without stipules. Stomata anomocytic. Flowers

solitary or in various types of cymose or racemose inﬂ o-

rescences, bisexual, actinomorphic or zygomorphic,

2-merous or less often 3-merous, usually entomophil-

ous. Sepals 2, sometimes 3 or 4, mostly caducous. Petals

usually twice as many as sepals and in 2 whorls, mostly

4, less often 6, sometimes 8–12 (up to 16), free or api-

cally connate or connivent, rarely absent (Macleaya and

Bocconia). Stamens numerous or 6–12, rarely only 4

(Hypecoum and Pteridophyllum), free or diadelphous

(Fumariaceae), when numerous developing in centripe-

tal sequence; anthers opening longitudinally. Tapetum

secretory or amoeboid. Microsporogenesis simultane-

ous. Pollen grains 2-celled, 3-colpate to pantocolpate

and pantoporate, seldom 2-colpate, rarely (Meconopsis,

Cathcartia) inaperturate. Gynoecium paracarpous, of 2

or less often 3–25 carpels; ovary superior, rarely almost

semi-inferior, mostly with numerous (up to 180 in

Glaucium) ovules, rarely two or only one (Bocconia,

Macleaya) ovule. Ovules anatropous, hemicampylotro-

pous or hemiamphitropous, or campylotropous, biteg-

mic, crassinucellate, with the micropyle formed by both

integuments. Female gametophyte monosporic of

Polygonum-type or rarely (Platystemon) tetrasporic of

Fritillaria-type. Endosperm nuclear. Fruits typically

capsular. Seeds small, arillate (except Hypecoaceae and

Pterydophyllaceae); seed coat formed by both integu-

ments; embryo minute and undifferentiated to linear and

elongate and more or less differentiated; endosperm

copious, oily. Producing various types of isoquinoline

alkaloids (including benzyl isoquinoline and aporphine

types and most notably protopine). Chromosomes large

to very small, n = 5–10, 14, 19, initially probably 6

(Safronova 1988a, b).

Close to Ranunculales and Berberidales (especially to

Hydrastidaceae and Podophyllaceae) and Glaucidiales

and shares a common origin with them. The Papaverales

differ from the other Ranunculanae mainly in their

paracarpous gynoecium with parietal placentation.

Key to Families

1 Plants producing milky or colored latex in articulated

laticifers or less often in elongate latex cells. Flowers

actinomorphic. Sepals 2 or 3, rarely 4, free or some-

times more or less connate, fully enclosing the bud

before anthesis. Petals usually in two cycles of two or

three, usually caducous, imbricate and often crum-

pled in bud, rarely lacking or in more than two cycles,

not spurred. Stamens mostly numerous, free.

Gynoecium of 2–25 carpels. Perennial or annual

herbs, rarely evergreen shrubs or small trees. Nodes

unilacunar. Leaves alternate, rarely opposite or ver-

ticillate, mostly variously divided, n = 5–10, 14

(Argemone), 19 (Romneya). . . . . . 1. papaveraceae

1 Plants without laticifers. Secretory idioblasts pres-

ent or absent. Flowers actinomorphic or zygomor-

phic. Sepals two, small, not enclosing the developing

bud. Petals four, spurred or not. Stamens four or six.

Gynoecium of two carpels.

2 Flowers nearly actinomorphic. Outer petals not

spurred. Stamens four, free.

3 Stemless perennial herbs with short thick

rhizomes. Leaves basal, oblanceolate, gradu-

ally narrowed below, pectinate, with 10–20

pairs of dentate segments, fern-like.

Inﬂ orescence an open thyrsus with small den-

tate bracts subtending cymules of 1–4 pedicel-

late ﬂ owers. Individual ﬂ owers small, white,

2- merous. Sepals two, small, caducous. Petals

four, entire, oblong, inner narrower than outer.

Stamens four, alternating the inner petals; ﬁ la-

ments short, not winged; anthers linear. Pollen

grains (2)3(4)-colpate, with perforate, sub-

echinulate tectum. Gynoecium with a long,

slender style and densely papillate biﬁ d stigma.

Ovules two or rarely four, anatropous to sub-

campylotropous. Fruits sili-culiform, 2-seeded

capsules dehiscing by two valves. Seeds ellip-

soidal, without elaiosomes, with small embryo,

n = 9. . . . . . . . . . . . . . 2. pteridophyllaceae

3 Annual herbs with slender stems. Leaves lan-

ceolate, 2–4 pinnatisect, the lobes linear to

narrowly obovate. Inﬂ orescence cymose,

many-ﬂ owered, with foliose bracts. Flowers

actinomorphic, 2-merous. Sepals 2, ovate to

lanceolate, herbaceous, with membranous,

often eroded margin. Petals four in two cycles,

small, outer petals entire to 3-lobed, inner pet-

als triﬁ d, with linear lateral lobes and stipitate,

Superorder RANUNCULANAE 95

cochleariform, ﬁ mbriate to denticulate median

lobe. Stamens four; ﬁ laments winged, basally

connate with the glandular tissue along the mar-

gins; anthers oblong. Pollen grains 2-colpate.

Gynoecium with two long and narrow stigmatic

branches. Ovules several, campylotropous. Fruit

a many-seeded lomentum or dehiscing by two

valves, n = 7, 8. . . . . . . . . . . . .3. hypecoaceae

2 Flowers bilaterally symmetrical or more often

transversely zygomorphic. One or both of the

outer petals usually basally saccate or spurred,

and the inner ones narrower and more or less

connate or sticky- connivent over the anthers.

Stamens six, diadelphous in a pair of lateral

phalanges usually with one or two nectary

glands at base. Fruits transversely septate elon-

gated or bladdery capsules or seldom 1-seeded

nuts. Perennial or annual herbs. Leaves alter-

nate or rarely (some Corydalis spp.) opposite,

pinnately to ternately divided. In some parts of

the plant, particularly below the leaf epidermis,

unbranched idioblasts present; their juice con-

tains alkaloids and fatty oils, and may be red-

dish or yellowish, but never milky, n = 8 or very

rarely 7. . . . . . . . . . . . . . . . . . 4. fumariaceae

1. PAPAVERACEAE

A. L. de Jussieu 1789 (including Chelidoniaceae

Martynov 1820, Eschscholziaceae Seringe 1847,

Platystemonaceae Lilja 1870). 26/250. Mostly temper-

ate and subtropical regions of the Northern Hemisphere,

especially the Mediterranean, western, central, and east-

ern Asia and southwestern parts of the USA; extending

to northern South America; 1 species of Papaver in

South Africa and 1 species in Cape Verde Islands.

1.1 CHELIDONIOIDEAE

Rhizomatous perennials or arborescent plants. Latex

orange, yellow or red. Hairs multicellular and termi-

nally uniseriate. Perianth 2-merous. Gynoecium

2-carpellate. Fruits opening by valves. Seeds mostly

arillate (except Glaucium and Dicranostigma). n = 5, 6,

7, 9, 10, chromosomes relatively large to very small. –

chelidonieae: Sanguinaria, Eomecon, Stylophorum,

Hylomecon, Coreanomecon, Chelidonium; glaucieae:

Glaucium, Dicranostigma; bocconieae: Macleaya,

Bocconia.

1.2 ESCHSCHOLZIOIDEAE

Perennial or annual herbs or arborescent plants.

Hairs unicellular. Perianth 2(rarely 3)-merous.

Gynoecium 2-carpellate. Fruits with ten conspicuous

longitudinal ribs, opening by valves and usually explo-

sively. Seeds nonarillate or (Dendromecon) arillate.

n = 6, 7, 8, chromosomes small (especially in

Hunnemannia), metacentric. – Dendromecon, Hunne-

mannia, Eschscholzia.

1.3 PAPAVEROIDEAE

Large suffruticose herbs to tiny annuals. Hairs multi-

cellular and multiseriate. Perianth 2- or 3-merous.

Gynoecium 2–24-carpellate. Stigma laterally conﬂ u-

ent. Fruits opening by three or usually more valves.

Seeds usually nonarillate (except Arctomecon). n = 7,

less often 6, 8 or (Argemone) 14 and (Romneya) 19,

chromosomes rather large or (Argemone) small, gener-

ally submetacentric and intercentric. – papavereae:

Papaver, Roemeria, Stylomecon, Meconopsis, Cath-

cartia, Argemone, Canbya; romnneyeae: Romneya,

Arctomecon.

1.4 PLATYSTEMONOIDEAE

Annual herbs with narrow and entire, often opposite or

verticillate leaves. Hairs multicellular and multi-

seriate. Perianth 3-merous. Gynoecium 3-carpellate or

(Platystemon) 6–25-carpellate. Carpel tips and stigmas

free. Fruits opening by basipetal separation of carpels

along placentas without formation of valves or

(Platystemon) breaking transversely into 1-seeded seg-

ments. n = 6, 7, 8, chromosomes metacentric, rather

large. – Platystemon, Hesperomecon, Meconella.

Chelidonioideae are the most archaic subfamily of

the Papaveraceae. However, they are rather heteroge-

neous and the Glaucieae differ in some respects from

the rest of the subfamily. According to Kadereit et al.

(1994), they differ by the possession of frondose bracts,

the absence of a seed appendage, and the absence of

chelidonin and stylopin as characteristic alkaloids of

the Chelidoniodeae s. str. Perhaps the best solution

will be the new subfamily Glaucioideae, as suggested

by Stevenson and Loconte (personal communication).

As regards Platystemonoideae, they represent simpli-

ﬁ ed rather than primitive forms (Ernst 1962, 1967).

They are close to the Papaveroideae, particularly to

Canbya (Ernst 1962: 1067; Kadereit et al. 1994), but

the valvular dehiscence within the carpel wall of the

fruit, which at maturity leaves behind the persistent

96 Subclass II. RANUNCULIDAE

placental bundles and the fused stigmatic mechanism,

clearly isolates Canbya from the Platystemonoideae

(Ernst 1967: 61). I agree with Ernst “that the similari-

ties between Canbya and Platystemonoideae are

manifestations of highly advanced features in

Papaveraceae and that Canbya probably can have

played no direct role in the evolution of

Platystemonoideae.”

2. PTERIDOPHYLLACEAE

Nakai ex Reveal and Hoogland 1991. 1/1. Japan

(Honshu).

Pteridophyllum.

Pteridophyllum racemosum is a rare plant that

grows in coniferous woods of central and northern

Honshu. It is usually included in the Papaveraceae

s. str., from which it differs markedly by the absence of

latex, fernlike pectinately pinnate leaves surrounded at

the base by round scales, and constantly 4-merous

androecium. Somewhat intermediate between

Papaveraceae and Hypecoaceae.

3. HYPECOACEAE

H. M. Wilkomm and J. M. C. Lange 1880. 1/18. From

the western Mediterranean to Mongolia and north-

western China.

Hypecoum.

Hypecoum is usually included in the Fumariaceae,

from which it differs by the morphology of corolla and

androecium as well as by embryological characters

(Iljina 1981) and seed coat anatomy (Iljina 1988). In

contrast to all other members of the Papaverales, in

Hypecoum the pollen is deposited not directly onto the

stigma but onto the central lobes of the inner petals,

which clasp the stamens and the style (Dahl 1989). It

differs from the Papaveraceae and Fumariaceae no less

than these two families from each other.

4. FUMARIACEAE

Berchtold et J. Presl 1820 (including Corydalaceae

Vest 1818). 18/530. Mostly northern temperate regions;

only a few are found south of the equator (a few species

of Corydalis on mountains in East Africa and the small

genera Phacocapnos, Cysticapnos, Trigono capnos,

and Discocapnos in southern Africa).

corydaleae: Dicentra, Capnoides, Adlumia,

Dactylicapnos, Corydalis, Roborowskia; fumarieae:

Pseudofu-maria, Sarcocapnos, Ceratocapnos,

Platy capnos, Fumari-ola, Fumaria, Cryptocapnos,

Rupicapnos, Discocapnos, Trigonocapnos, Phaco-

capnos, Cysticapnos.

Close to the Papaveraceae, especially to the

Chelidonioideae and Eschscholzioideae, with which

they share a common origin.

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Order 28. GLAUCIDIALES

Perennial herbs with well-developed and creeping rhi-

zome producing a single long-petiolate leaf on its end.

Stem solitary or sometimes two together, simple, rather

stout, minutely poberulous, eventually glabrescent

above. Vascular bundles amphicribral; xylem barely

concave at the outer edge, but not V-shaped. Vessels in

the rhizome mostly with simple perforations, but some-

times with scalariform or, rarely, reticulate perforations;

lateral pitting scalariform. Fibers with bordered pits.

Nodes multilacunar. Leaves few, alternate, apparently

arranged in 1/2 divergence, simple, palmately lobed,

without stipules; basal leaves long-petiolate, cauline

leaves usually two, rarely three, borne on the uppermost

part of the stem, reniform or cordate-orbicular; verna-

tion conduplicate. Stomata anomocytic. Flowers large,

solitary and terminal, bisexual, spirocyclic, actinomor-

phic, apetalous. Bract foliaceous, sessile, reniform to

suborbicular, cordate, sharply incised-dentate. Sepals

four, large, decussate, broadly obovate-rhomboid, peta-

loid, spreading, caducous. Stamens very numerous, spi-

rally arranged, fasciculate, initiated centrifugally;

ﬁ laments ﬁ liform, slightly enlarged and abruptly con-

stricted below the anther; anthers tetrasporangiate,

opening longitudinally. Pollen grains 2-celled, 3-colpate,

with punctate/spinulose tectum in which spinules are

more numerous. Carpels usually two (rarely solitary

or 3), shortly connate at the base by the intervention of

receptacular tissue, conduplicate, with a bipartite stigma

on a very short stylodium and with distinct furrows both

along the ventral suture and along the dorsal side, with

15–30 ovules in each carpel. Ovules, anatropous, biteg-

mic, pseudo-crassinucellate; the outer integument vas-

cularized. Endosperm nuclear. Female gametophyte of

Polygonum-type. Fruiting carpels laterally compressed,

subquadrate, dehiscing both ventricidally and dorsicid-

ally. Seeds pendulous in two rows from upper edge of

fruiting carpels, the members of the two rows strictly

alternating but forming a single line, compressed,

broadly winged, obovate, about 1.5 cm long, brown,

exotestal, with minute embryo and copious endosperm.

Cotyledons have a tendency to fuse. Producing glau-

palol, a substance with a coumarin skeleton. No alka-

loids is detected, n = 10, chromosomes small and short.

Glaucidiaceae deﬁ nitely belong to Ranunculidae, but

from all of them differ by the assemblage of characters.

The nearest are probably the Ranunculaceae or maybe

the Hydrastidaceae (see Tobe 2003) but from them they

Superorder RANUNCULANAE 99

differ in conduplicate leaf vernation, vascular bundles

(as in Hydrastidaceae), scalariform lateral pitting (as in

Hydrastidaceae), vascular anatomy of the ﬂ ower, mor-

phology of the gynoecium, micropyle formation, vascu-

larized outer integument, the absence of nectar discs,

fasciculate stamens, initiated centrifugally (as in

Paeoniaceae), pollen morphology, fruiting carpels

dehiscining both ventricidally and dorsicidally, broadly

winged seeds, absence of alkaloids and presence of glau-

palol, a substance with a coumarin skeleton, the basic

number of chromosomes (see Tobe and Keating 1985).

1. GLAUCIDIACEAE

Tamura 1972. 1/1. High mountains of central and

northern Japan (Hokkaido and northern and central

districts of Honshu).

Glaucidium.

Bibliography

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genus. Bot. Mag. (Tokyo) 94: 207–224.

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Order 29. PAEONIALES

Perennial herbs or less often small shrubs or subshrubs

with scattered secretory cells in the parenchymatous

tissues of the stems. Rhizome and roots thickened,

napiform or bulbous. Vascular bundles amphicribral (as

in Hydrastidaceae and Glaucidiaceae). Vessels very

small, angular, with scalariform or simple perforations

that have 1-3(-7) bars (in metaxylem 4–12 bars); lateral

pitting alternate. Fibers with bordered pits. Rays hetero-

cellular to homocellular. Wood parenchyma diffuse.

Nodes pentalacunar with ﬁ ve traces. Leaves large,

alternate, ternately or ternate-pinnately compound or

dissected; petioles dilated at base; leaﬂ ets entire,

3–5-lobed to – parted, or dissected into linear segments,

cuneate to rounded at base; stipules absent. Stomata

anomocytic. Flowers large, terminal, solitary or some-

times a few together, spiral or spirocyclic, bisexual, acti-

nomorphic. Sepals (3-)5(-7), persistent. Petals

morphologically equivalent to sepals, mostly 5, less

often up to 10 (rarely up to 13), large, free. Stamens

numerous, with thin ﬁ lament united into 5 fascicles, the

members of which almost always develop in centrifugal

sequence (except Paeonia japonica, Sawada 1971);

anthers tetrasporangiate, extrorse, opening longitudi-

nally. Tapetum secretory. Microsporogenesis simultane-

ous. Pollen grains 2-celled, 3-colpate/colporoidate/

colporate, with ﬁ nely punctate and rugulose or perforate

tectum. Gynoecium apocarpous, of (2-)5, rarely up to

8 and even to 15 carpels; carpels coarse and thick-walled,

with almost sessile expanded stigma; gynoecium sur-

rounded by a ﬂ eshy, lobed nectary disc that is probably

a modiﬁ ed part of the androecium (Eames 1953, 1961).

Ovules several or many in each carpel, anatropous,

bitegmic (with massive, slightly vascularized outer

integument, which is longer than the inner one), crassi-

nucellate, with nucellar cup. Female gametophyte of

Polygonum-type. Endosperm nuclear. Embryogenesis of

unique type: the zygote nucleus undergoes repeated free

nuclear divisions to form a coenocyte with the nuclei

restricted to a peripheral layer around a large central

vacuole; wall formation proceeds centripetally and the

proembryo becomes cellular. Fruits of follicles. Seeds

medium-sized to large, with more or less well-developed

dark red aril; seed coat formed by the outer integument

(testal); embryo minute, differentiated; endosperm

copious. Producing phenolic substance and glycosides,

such as paeonol, paeonoside, paeoniﬂ orin, etc., ethereal

100 Subclass II. RANUNCULIDAE

oils, ﬂ avones, seed polysaccharides (amyloids) and

alkaloids nearly lacking; n = 5, somewhat n = 10.

The Paeoniales are closely related to the Glaucidiales

(Melville 1982), from which they differ by tricolporate

pollen grains, persistent and nondecussate sepals, not

sharply limited perianth (the phyllotactic spiral in

Paeonia is continuous from leaves through bracts, sepals,

petals, stamen trunks, and carpels, Eames 1961: 433),

somewhat convex receptacle, lobed nectary disc sur-

rounding the gynoecium, the longer outer integument,

arillate seeds, presence of ethereal cells in parenchyma-

tous tissues, long chromosomes, absence of glaupalol,

and presence of amyloids in seeds. Both orders share a

common origin from some ranunculalean ancestor.

Afﬁ nities with the Dilleniaceae seem less probable.

1. PAEONIACEAE

Raﬁ nesque 1815. 1/35–40. North Africa (north of

Sahara), Europe, subtropical, temperate, and partly cold

regions of Asia, western regions of North America.

Paeonia.

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