Takhtajan Armen. Flowering Plants

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162 Subclass IV. CARYOPHYLLIDAE

sessile glands), or not heterophyllous. Leaves

and stems are beset with peltate hairs and with

characteristic multicellular, stalked, or sessile

glands secreting a sticky mucilage that traps

insects. Interxylar (included) phloem present.

Vessel elements extremely large (except in

Habropetalum); lateral pitting alternate. Fibers

with large, bordered pits. Axial parenchyma

paratracheal (vasicentric) and apotracheal.

Vasicentric tracheids present. Leaves entire or

crenulate, estipulate, pinnately veined; the mid-

rib usually prolonged and forked into two

recurved hooks or pigtailed tendrils. Stomata

actinocytic or encyclocytic. Flowers of moder-

ate size, in a lax, axillary or supra-axillary,

cymose inﬂ orescences, bracteate. Sepals ﬁ ve,

small, free or connate at the base into a short

tube, valvate or open in bud, persistent. Petals

ﬁ ve, free, contorted, caducous. Stamens 10(-

30), free or slightly connate; anthers ovoid or

oblong, basiﬁ xed, introrse, appendaged, open-

ing longitudinally. Pollen grains 3(-4)- colporate.

Gynoecium of two or ﬁ ve united carpels; sty-

lodia either two, free, ﬁ liform, with capitate

stigmas, or ﬁ ve, free, ﬁ liform, with minute stig-

mas, or ﬁ ve, slightly connate at the base and

with plumose stigmas. Ovary superior, 1-locular,

with numerous ovules on parietal placentas.

Ovules anatropous, bitegmic, crassinucellate

(but according to Schmid 1964, ovules in

Triphyophyllum semitenuinucellate). The outer

integument forms a characteristic cover.

Endosperm probably nuclear. In connection

with the development of the integumentary

cover, the female gametophyte (and therefore

also the endosperm) elongate perpendicularly.

Fruits loculicidal capsules of two or ﬁ ve valves,

spreading widely, opening well before maturity,

and bearing the few seeds peltately attached on

greatly elongate, thickened, rigid funicles. Seeds

large, discoid, either thick and surrounded by

narrower wing, or thin and surrounded by broad

satiny wing; embryo large, discoid-obconic,

axile, mostly surrounded by the copious

starchy endosperm. Produce naphthoquinones

like plumbagin, and naphthylisoquinoline alka-

loids, e.g. dioncophylline (Bringmann and

Pokorny 1995; Porembski and Barthlott 2002),

and ellagic acid, n = 12, 18 (Triphyophyllum

peltatum).. . . . . . . . . . . 2. dioncophyllaceae.

2 Flowers unisexual, apetalous. Leaves transformed

into pitchers. Herbaceous or sometimes more or

less woody plants with a small amount of second-

ary growth, erect, prostrate, or often climbing by

aid of the leaf tendrils, terrestrial or epiphytic.

Various kinds of multicellular glands and peltate

hydathodes are widely distributed on the stem

and leaves. Schmid-Hollinger (1971) recognises

ﬁ ve types of non-glandular trichomes: multicel-

lular simple hairs, hairs with teeth, tufted, rosette

and arachnoid hairs. Cortical bundles occur in

young stem, and medullary vascular bundles in

the stem of certain species. Characteristic spiral

cells occur in the parenchymatous tissues of both

stem and leaf. Vessels dimorphic, including wide,

short elements along with fusiform vessel mem-

bers, both usually with simple perforation, true

tracheids present. Rays mostly biseriate or unise-

riate; multiseriate rays more than two cells wide

are relatively scarce. Axial parenchyma banded

and diffuse. Leaves alternate, simple, estipulate,

when fully developed composed of a more or less

distinct petiole, a winged or expanded portion

(blade) followed by a stout and often coiled ten-

dril (a prolongation of the leaf midrib), usually

terminated by a large, pendant, often brightly col-

ored, urceolate or cylindrical pitcher (ascidium)

that has a recurved rim and a ﬂ attened lid (oper-

culum) projecting over the mouth, but not closing

it except in the young state. Below the base of the

lid on the outside of the pitcher is a spur (an

appendage); at the entrance of the pitcher are

numerous nectar-glands, below which the interior

is slippery, and for some distance below are diges-

tive glands. The second appendage, if developed,

also lies on the midline, but at the apex of the lid;

the apical appendage is more ﬁ liform, and may

not always be glandular (Cheek and Jebb 2001).

Stomata anomocytic. Flowers small, in racemes

or panicles, dioecious. Sepals usually four, rarely

three, imbricate, free or less often basally con-

nate, glandular and nectariferous within. Stamens

(2-)8–25, rarely 4–6; ﬁ laments united into a col-

umn; anthers crowded but distinct, 2-locular,

extrorse, tetrasporangiate, opening longitudinally

by slits. Pollen grains 3-celled, in tetrads, spinu-

lose, with indistinct apertures. Gynoecium syn-

carpous, of four or very rarely three or six carpels

opposite the sepals; ovary superior, 4- or rarely

3-locular, with numerous ovules multiseriate on

Superorder NEPENTHANAE 163

axile placentas; stigma discoid, on a very short

style or sessile. Ovules numerous, multi seriate on

axile or laminar placentas, ascending, anatropous,

bitegmic, crassinucellate. Endosperm nuclear.

Fruits elongated, sometimes stipitate, leathery

loculicidal capsules. Seeds numerous (100–500),

mostly ﬁ liform, ascending, imbricate, often

winged (usually, with a narrow or hair-like wing

at either end), or wingless; seed coat formed by

both the integuments, but mainly exotestal;

exotesta with much thickened inner walls. Embryo

minute but well-differentiated, straight, cylindri-

cal, surrounded by starchy, proteinaceous and

oily endosperm, sometimes endosperm absent.

Tanniniferous, producing acetogenic napthoqui-

none plumbagin, cyanidin, and also kaempferol

and quercetin, n = 40. . . . . . . 3. nepenthaceae.

1 Ovary inferior or partly inferior. Sympodially

branched, scandent shrubs or lianas, with hooked or

twining shoot ending of each member of the sympo-

dium, “with juvenile growth often heteromorphic,

erect, shrubby, monopodial, unbranched or sparsely

branched, and without hooks, with adult growth

comprising elongated, usually little-branched stems

that bear smaller extra-axillary latral stems” (Taylor

et al. 2005). Plant externally glabrous, often with

calcium oxalate crystals. Primary cortex contains

isolated, thick-walled secretory cells. Vessels with

simple perforations in very oblique end walls. Sieve-

element plastids Ss-type. Fibers with large, evidently

bordered pits. Rays homogeneous. Axial paren-

chyma commonly in tangential bands. Nodes trila-

cunar with three traces. Leaves pinnately veined,

with very small, mostly caducous stipules, or estipu-

late, and beset with minute, embedded, peltate, mul-

ticellular, waxy glands. Stomata actinocytic

(surrounded by more or less distinct subsidiary

cells). Flowers small, in panicles, in spikes, and in

cymes; the terminal inﬂ orescence unit often appar-

ently cymose (axillary or terminal), with strongly

articulated pedicels, 5-merous, rarely (Ancistrocladus

grandiﬂ orus) 4-merous. Sepals free to connate into

a short tube, at length adnate to the base of the ovary;

lobes quincuincial, imbricate, becoming unequally

enlarged and winglike in fruit. Petals more or less

ﬂ eshy, shortly connate or cohering, imbricate.

Stamens (8-)10 in one cycle or rarely 5 (A hamatus)

or 15 (A. robertsoniorum); ﬁ laments short, ﬂ eshy,

connate below and adnate to the bases of the petals;

anthers basiﬁ xed, introrse or latrorse, tetrasporan-

giate, opening longitudinally. Pollen grains 2(4)-col-

porate. Gynoecium of three united carpels; ovary

1-locular, with one basilateral ovule on a short

funiculus; stylodia free or connate, articulated at the

base and thickened upward, with more or less hip-

pocrepiform or puntoﬁ d stigmas. Ovules ascending,

hemitropous, bitegmic, on a short funiculus, crassi-

nucellate. Endosperm cellular. Fruits dry, woody,

indehiscent, surrounded by a corky hypanthium and

crowned by spreading or erect winglike calyx-lobes,

ﬂ oating in water. Seeds large, with a thin, membra-

nous seed coat, strongly ruminate, hard, starchy,

sometimes ruminate endosperm, with testa intrud-

ing between folds, and fairly small, straight embryo.

Produce acetogenic isoquinoline alkaloids, cyanidin

and delphinidin; kaempferol, quercetin, myricetin

and ellagic acid. . . . . . . . . 1. ancistrocladaceae

1. ANCISTROCLADACEAE

Planchon ex Walpers 1851. 1/16. Tropical Africa,

southern Asia, and from eastern Himalayas to West

Malesia.

Ancistrocladus.

Numerous molecular studies (particularly Meimberg

et al. 2000), provide evidence for the sister

group relationship between Ancistrocladaceae and

Dioncophyllaceae and their close relationship with

Nepenthaceae, Drosophyllaceae, and Droseraceae,

conﬁ rming the earlier view of Airy Shaw (1952).

2. DIONCOPHYLLACEAE

Airy Shaw 1952. 3/3. Western tropical Africa.

Dioncophyllum, Triphyophyllum, Habropetalum.

Closely related to the Ancistrocladaceae and Droser-

aceae. The traps on young stems and leaves, constituted

by multicellular, stalked or sessile secretory glands,

recalling those of Ancistrocladaceae and Droseraceae,

which secrete sticky, acid mucilage. In their alkaloids

Dion cophyllaceae strongly resemble Ancistrocladaceae

(Hegnauer 1987; Bringmann et al. 1991).

3. NEPENTHACEAE

Berchtold et J. Presl 1820. 1/87. Madagascar (2),

Seychelles (1), Sri Lanka (1), northeastern India

164 Subclass IV. CARYOPHYLLIDAE

(1, Assam), Southeast Asia (6–9), northern Australia

(1), New Caledonia (1), but mostly in Malesia (82).

Nepenthes (including Anurosperma).

Related to Ancistrocladaceae and Dioncophyl-

laceae.

4. DROSOPHYLLACEAE

Chrtek, Slaviková et Studnicka 1989. 1/1. Portugal,

southern Spain, Morocco.

Drosophyllum.

For a long time Drosophyllum is included in

Droseraceae, but recent studies have indicated an iso-

lation of this genus within Droseraceae. Metcalfe

(1952) showed that Drosophyllum shares the stalked,

non-irritable glands, and the whole trapping mecha-

nism, with Triphyophyllum of Dioncophyllaceae.

Strong evidence for the afﬁ nity of Drosophyllum with

other members of the Nepenthales has been provided

by cladistic analysis of matK gene sequences

(Meimberg et al. 2000).

5. DROSERACEAE

R. A. Salisbury 1808 (including Aldrovandaceae

Nakai 1949, Dionaeaceae Raﬁ nesque 1837). 3/112.

Drosera (110) subcosmopolitan with concentration in

Australia and New Zealand, Dionaea (1) in southeastern

United States, and Aldrovanda (1) throughout much of

Eurasia – extends south to northeastern Australia.

Drosera, Dionaea, Aldrovanda.

Probably related to the Nepenthaceae, which is sup-

ported also by molecular data (Albert et al. 1992).

Meimberg et al. (2000) placed Droseraceae sister to a

clade comprising Dioncophyllaceae, Ancistrocla-

daceae and Nepenthaceae.

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A. Takhtajan, Flowering Plants, 167

Trees, shrubs, and herbs. Ethereal oil cells wanting.

Vessels with scalariform or simple perforations. Sieve-

element plastids of S-type or rarely of P-type. Nodes

tri-multilacunar or unilacunar. Leaves simple or less

often compound, with or without stipules. Stomata of

various types, mostly anomocytic. Flowers usually

with double perianth, mostly bisexual, spiral, spirocy-

clic or cyclic. Petals free or less often corolla sympeta-

lous. Pollen grains 2-celled or seldom 3-celled,

3-colpate or of tricolpate-derived type. Gynoecium

apocarpous or more often carpels variously united;

stylodia distinct or more or less connate; ovary

superior or inferior. Ovules mostly bitegmic or less

often unitegmic, commonly crassinucellate. Endosperm

nuclear or less often cellular. Fruits of various types.

Seeds mostly with endosperm.

Dilleniidae are one of the largest subclasses of the

magnoliopsids and one of their central groups, archaic

members of which have many features in common

with the Magnoliidae.

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Superorder DILLENIANAE

Order 48. DILLENIALES

Mostly trees (sometimes buttresses, or tortuous to

small rosette trees), and shrubs, rarely woody lianas,

subshrubs (Pachynema), or semiherbaceous from a

woody rhizome (Acrotrema); raphides common in

wood. Vessel elements angular to circular, from

medium to extremely long, usually very long with sca-

lariform perforations that have 5–130 bars (mostly

15–40) or scalariform and simple perforations; lateral

pitting from scalariform to opposite. Fibers with dis-

tinctly bordered pits. Rays heterogeneous. Axial paren-

chyma apotracheal diffuse and aggregate diffuse, often

paratracheal diffuse to incomplete vasicentric. Sieve-

element plastids of S-type. Nodes trilacunar, pentala-

cunar, multilacunar, or sometimes unilacunar with one

trace per gap. Leaves usually alternate, very rarely

opposite, entire or serrate to dentate, rarely pinnatiﬁ d

or trilobed, pinnately veined, often with prominent

parallel lateral veins, seldom much reduced and needle-

like (species of Hibbertia) or scalelike (species of

Pachynema with stems and branches ﬂ attened into

phylloclades). Stipules usually wanting, but rarely pet-

ioles with stipulelike, often wholly or partly caducous

wings. Walls of epidermal cells mostly siliciﬁ ed (as in

Magnoliaceae and Calycanthaceae). Stomata of vari-

ous types, but mostly anomocytic. Flowers small to

medium-sized, rarely large, in various cymose or rac-

emose inﬂ orescences or sometimes solitary, bisexual

or rarely unisexual (monoecious or dioecious),

Subclass V. DILLENIIDAE

168 Subclass V. DILLENIIDAE

actinomorphic or sometimes zygomorphic (but only

with reference to the androecium), with spirocyclic

perianth. Receptacle ﬂ at or conical between the carpels

(Dillenia). Sepals mostly ﬁ ve, rarely more (up to 15)

or less (3), strongly imbricate, persistent. Petals ﬁ ve

or rarely fewer, imbricate, often crumpled in bud,

deciduous. Stamens usually numerous (from 50 to 500

in Tetracera and to 900 in Dillenia ovalifolia), rarely

10 or fewer. Filaments usually filiform, rarely broad

and ﬂ attened (Pachynema), variously united into

fascicles at the base; the members of each fascicle

originating in centrifugal sequence, or the whole

androecium developing centrifugally. The androecium

of Schumacheria is unique in its placement at one

side of the gynoecium with the large number of

stamens united basally by their ﬁ laments into an oblique

cylindric column. Anthers tetrasporangiate, often more

or less embedded in the connective tissue, from short

to very elongate, sometimes with more or less prolonged

connective, opening by longitudinal slits or apical pores.

Tapetum secretory. Microsporogenesis simultaneous.

Pollen grains 2-celled, 3-colpate (Acroterma, Dillenia,

Hibbertia, Pachynema), 4-colpate or 3-colporate,

tectate-columellate, with an incomplete tectum, ﬁ nely

lunctate to coarsely reticulate. Gynoecium usually of

1–20 carpels, mostly of several carpels; carpels free,

less often partly connate at the base, rarely fused to

above the middle (Curatella) or even to the apex

(Neowormia); carpels more or less conduplicate and

sometimes not fully sealed. Stylodia usually slender

and elongate, with an apical, often capitate, minute

stigma. Ovules one to many in each carpel, apotropous,

anatropous to amphitropous or campylotropous, biteg-

mic, crassinucellate; micropyle zigzag or exostomal.

Female gametophyte of Polygonum-type. Endosperm

nuclear. Fruits apocarpous or syncarpous, mostly dry

and dehiscent, sometimes indehiscent (Dillenia). Seeds

mostly with a well-developed crested or laciniate

funicular aril (very large in Dellenia sarmentosa

and Tetracera spp.), less often with the aril vestigial

or even wanting. Seed coat formed by both integu-

ments and with sclerotic endotesta and tracheidal

exotegmen (endotestal-exotegmal); embryo usually

minute, straight, more or less differentiated; endosperm

copious, oily, and proteinaceous, sometimes trumi-

nate. Tanniniferous, containing a great diversity of

ﬂ avonoids, but mostly without alkaloids and always

without benzylisoquinoline alkaloids, n = 4, 5,

8–10, 12, 13.

The Dilleniaceae are a family of considerable

morphological and ecological diversity (Hutchinson

1969; Dickison 1970b, 1982; Stebbins 1974;

Cronquist 1981). They are also very heterobathmic.

Although they retain some primitive features in wood

anatomy and ﬂ ower morphology (apocarpous gynoe-

cium, more or less conduplicate carpels and stamens

that sometimes have a prolonged connective),

they have also many features of specialization,

including specialized androecium and specialized

stylodia with apical non-decurrent stigma. Although

the Dilleniaceae are the most archaic family in the

Dilleniidae, they have already lost direct links with

the Magnoliidae. However, they occupy an interme-

diate position between the Magnolianae and the

Ericanae and Violanae.

1. DILLENIACEAE

R.A. Salisbury 1807 (including Hibbertiaceae J. Agardh

1858, Soramiaceae Martynov 1820). 12/400. Almost

pantropical, but centered in Asia and Australasia,

especially in Australia. Comparatively rare in Africa,

where it is represented only by members of the pan-

tropical genus Tetracera. Hibbertia is native to both

Australasia and Madagascar.

1.1 DILLENIOIDEAE

Perforations exclusively scalariform; lateral pitting

scalariform-transitional. Anthers linear or oblong,

loculi very rarely slightly divergent. – dillenieae:

Dillenia, Neowormia, Schumacheria, Didesmandra,

Acrotrema; hibbertieae: Hibbertia, Pachynema.

1.2 DELIMOIDEAE (TETRACEROIDEAE)

Perforations scalariform and simple; lateral pitting

opposite-transitional. Anthers short, loculi divergent,

separated at the base by the thickened connective. –

Tetreacera, Curatella, Pinzona, Doliocarpus, Davilla.

Bibliography

Baretta-Kuipers T. 1972. Some remarks on the wood structure of

Pinzona and allied genera of the subfamily Tetraceroideae

(Dilleniaceae). Acta Bot. Neerl. 21: 573–577.

Dickison WC. 1967a. Comparative morphological studies in

Dilleniaceae: I. Wood anatomy; II. The pollen. J. Arnold

Arbor. 48: 1–29, 231–240.

Superorder ERICANAE 169

Dickison WC. 1968. Comparative morphological studies in

Dilleniaceae: III. The carpels. J. Arnold Arbor. 49:

317–329.

Dickison WC. 1969. Comparative morphological studies in

Dilleniaceae: IV. Anatomy of the node and vascularization of

the leaf. J. Arnold Arbor. 50: 384–400.

Dickison WC. 1970. Comparative morphological studies in

Dilleniaceae: V. Leaf anatomy; VI. Stamens and young stem.

J. Arnold Arbor. 51: 89–113, 403–418.

Dickison WC. 1971. Comparative morphological studies in

Dilleniaceae. VII. Additional notes on Acrotrema. J. Arnold

Arbor. 52: 319–333.

Dickison WC. 1979. A note on the wood anatomy of Dillenia

(Dilleniaceae). IAWA Bull. 2/3: 57–60.

Dickison WC, JW Nowicke, and JJ Skwarla. 1982. Pollen mor-

phology of the Dilleniaceae and Actinidiaceae. Am. J. Bot.

69: 1055–1073.

Dulci Elizabeth J and KK Lakshmanan. 1984. Further contribu-

tions to the embryology of Dillenia suffruticosa (Griff.)

Martelli. J. Indian Bot. Soc. 63: 353–359.

Endress PK. 1997. Relationships between ﬂ oral organization,

architecture, and pollination made in Dillenia (Dilleniaceae).

Plant Syst. Evol. 206: 99–118.

Gurni AA and K Kubitzki. 1981. Flavonoid chemistry and

systematics of the Dilleniaceae. Biochem. Syst. Ecol. 9:

109–114.

Horn JW. 2002. Phylogenetics of the Dilleniaceae. In Botany

2002: Botany in the Curriculum, Abstracts: 128, Madison.

Horn JW. 2007. Dilleniaceae. In: K Kubitzki, ed. The families

and genera of vascular plants, vol. 9, pp. 132–154. Springer,

Berlin/Heidelberg/New York.

Kubitzki K. 1968. Flavonoide und Systematik der Dil-leniaceen.

Ber. Deutsch. Bot. Ges. 81: 238–251.

Sastri RLN. 1958. Floral morphology and embryology of some

Dilleniaceae. Bot. Not. 111: 495–511.

Swamy BGL and K Periasamy. 1955. Contributions to the

embryology of Acrotrema arnottianum. Phytomorphology 5:

301–314.

Vyshenskaya TD and GG Oganezova. 1991. Dilleniaceae.

In: A Takhtajan, ed. Comparative seed anatomy, vol. 3,

pp. 163–171. Nauka, Leningrad (in Russian).

Wilson CL. 1965. The ﬂ oral anatomy of the Dilleniaceae: I.

Hibbertia Andr. Phytomorphology 15: 248–274.

Wilson CL. 1973 (1974). The ﬂ oral anatomy of the Dilleniaceae:

II. Genera other than Hibbertia. Phytomorphology 23:

35–42.

Superorder ERICANAE

Order 49. PARACRYPHIALES

Small to moderate-sized trees with unicellular,

unbranched hairs. Styloid crystals of calcium oxalate

present in the bark, pith, and xylem and phloem paren-

chyma. Vessel elements extremely long, angular to

rounded in transverse section; perforations scalariform

in almost vertical end walls, with numerous (up to 203)

bars; lateral pitting scalariform to transitional. Fibers

thick-walled, with numerous bordered pits. Rays

heterogeneous, of very primitive type. Axial parenchyma

diffuse and diffuse-in-aggregates, apotracheal as well

as scanty paratracheal. Phloem of primitive type. Sieve-

elements of S-type. Nodes trilacunar. Leaves subver-

ricillate, simple, ﬁ nely serrate, pinnately veined, teeth

vascularized by a single medial vein extending to the

tooth apex, leathery, estipulate. Stomata anomocytic.

Flowers sessile, in terminal, paniclelike, compound

inﬂ orescences consisting of many spikes, bisexual or

occasionally exclusively male (without pistillodia) and

borne at the lower levels on the inﬂ orescence axis.

Perianth consists of four caducous, decussate, concave,

free segments (sepals?); aestivation of the perianth parts

cochleate, having one large helmet-shaped segment

that encloses or covers the others. Stamens usually

8 (–11), in a single cycle, free; ﬁ laments of the male

ﬂ owers conspicuously swollen, ﬁ laments of bisexual

ﬂ owers sometimes accrescent. Anthers tetrasporan-

giate, basiﬁ xed, opening longitudinally. Pollen grains

3-colporate, tectate-columellate, reticulate or occasion-

ally transitional between reticulate and rugulate.

Gynoecium of 8–15, laterally connate, conduplicate

carpels that are also ventrally adnate to the solid column

of central tissue. Stigmas sessile and conduplicately

folded; ovary superior, 8–15-locular, with four rather

small ovules in each locule borne in a single row on

axile placentas. Ovules anatropous, unitegmic, crassi-

nucellate. Endosperm cellular. Fruits capsules with

mature carpels separating from the central column

(except for a distal connection maintained by two

strands apparently representing the ligniﬁ ed ventral

carpellary vascular bundles, Dickison and Baas 1977),

spreading out from the base and opening ventrally.

Seeds small, compressed, winged, exarillate, exotestal,

with sinuous anticlinal walls, inner walls ligniﬁ ed;

embryo straight, well differentiated; endosperm copious.

As Dickison and Baas (1977) point out, Paracryphia

is distinguished by an unusual combination of

characters. “It is remarkable that most characters

from vege tative anatomy are distinctly primitive

or at least un-specialized, whereas those of repro-

ductive organs can be regarded as advanced or special-

ized.” Paracryphia was originally included into the

Eucryphiaceae but was later separated as its own

family by Airy Shaw (1965). Dahlgren (1983) placed

Paracryphiaceae in his suborder Theineae of the order

170 Subclass V. DILLENIIDAE

Theales, whereas according to Cronquist (1981: 336)

Paracryphia is reasonably at home in the Theales. He

mentions that the persistent central column is a common

(though by no means consistent) feature of capsular

fruits in the Theales, and the manner of dehiscence in

Paracryphia is reminiscent of Medusagyne. In my

opinion Paracryphiaceae are distinct enough to be

entitled to the status of an order (Takhtajan 1987, 1997).

Paracryphiales have many features in common with

the Dilleniales, Theales, and Ericales (see Dickison

and Baas 1977; Schmid 1978). This includes sieve-

element plastids that are extremely similar in size and

starch grain characters to those of Dilleniaceae,

Theaceae, and Actinidiaceae, and are almost identical

to the plastids of Hibbertia in the Dilleniaceae (Behnke

1985). Paracryphiales are possibly an early divergence

from an ancient dillenialean stock and one of the evo-

lutionary blind ends. Paracryphia is one of the most

remarkable ancient relicts.

1. PARACRYPHIACEAE

Airy Shaw 1965. 1/1. New Caledonia.

Paracryphia.

Bibliography

Agababian VS and EL Zavarian. 1971. Palynotaxonomy of the

genus Paracryphia Bak. f. Biol. Zhurn. Armenii 24 (12):

35–40 (in Russian).

Airy Shaw HK. 1965. Diagnoses of new families, new names,

etc., for the seventh edition of Willis’s Dictionary. Kew Bull.

18: 249–273.

Dickison WC and P Baas. 1977. The morphology and relation-

ships of Paracryphia (Paracryphiaceae). Blumea 23:

417–438.

Melikian AP. 1991. Paracryphiaceae. In: A Takhtajan, ed.

Comparative seed anatomy, vol. 3, p. 175. Nauka, Leningrad

(in Russian).

Schmid R. 1978. Actinidiaceae, Davidiaceae, and

Paracryphiaceae: systematic considerations. Bot. Jahrb. Syst.

100: 196–204.

Steenis CGGJ van. 1950. Note on Paracryphia Baker

(Eucryphiaceae). Bull. Jard. Bot. Buitenzorg 18: 459–460.

Order 50. THEALES

Trees, shrubs, or sometimes woody lianas or herbs.

Often with secretory cells, cavities, or channels. Vessel

elements from short to very long, with scalariform or

simple perforations; lateral pitting from scalariform to

alternate. Fibers with bordered or less often simple

pores. Rays mostly heterogeneous. Axial parenchyma

mostly apotracheal. Young stems often with stratiﬁ ed

phloem. Sieve-element plastids of S-type. Nodes unila-

cunar, less often pentalacunar (Oncothecaceae), or

penta- to multilacunar (Caryocaraceae). Leaves alter-

nate or opposite, simple or seldom compound, mostly

entire, estipulate, or very rarely (Caryocaraceae) stipu-

late. Stomata of various types, mostly paracytic. Flowers

of medium size or often large, axillary and solitary or in

various kinds of inﬂ orescences, usually bisexual, mostly

actinomorphic. Perianth spiral or spirocyclic (many

Theaceae) or more often cyclic, mostly 5-merous.

Calyx often subtended by 2-several bracteoles, these

sometimes gradually passing into sepals. Sepals imbri-

cate, free or often basally connate. Petals imbricate or

sometimes contorted, free or rarely connate at the base,

rarely (Anthodiscus in the Caryocaraceae) connate

above to form a calyptra, or corolla sympetalous

(Oncothecaceae). Stamens numerous to 5-4, free or

connate into bundles, when numerous developing in

centrifugal sequence. Anthers tetrasporangiate, basi-

ﬁ xed or dorsiﬁ xed, less often versatile, opening longitu-

dinally or rarely (spp. of Eurya and Pentaphylacaceae)

by short, apical, porelike slits; connective sometimes

more or less prolonged (very long in Anneslea frapans

of the Theaceae). Tapeturn secretory. Microsporogenesis

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

Gynoecium of 2-many variously united carpels.

Stylodia free or more or less connate; ovary superior,

rarely semi-inferior or inferior, with as many locules as

carpels or unilocular, usually with many ovules on axile

placentas or on intruded parietal placentas. Ovules ana-

tropous or less often hemitropous or more or less

campylotropous (Pentaphylacaceae), apotropous, biteg-

mic or (Oncothecaceae) unitegmic, crassinucellate

(Pentaphylacaceae) or more often tenuinucellate.

Female gametophyte of Polygonum- or Allium-types,

rarely Adoxa-type. Endosperm nuclear. Fruits of vari-

ous types. Seeds exarillate, seed coat formed mainly or

only by the outer integument; typically mesotestal with

sclerotic cells; endosperm copious or scanty, often very

reduced or absent; embryo small to large, straight or

curved.

The order Theales is evidently derived from early

dillenialean stock. It differs from the Dilleniales mainly

in the syncarpous gynoecium and mesotestal and

mostly exarillate seeds.

Superorder ERICANAE 171

Key to Families

1 Leaves simple. Nodes unilacunar or seldom trilacu-

nar or pentalacunar.

2 Flowers in axillary, pendulous racemes or

spikes, 4-merous throughout, closely sub-

tended by two well-developed bracteoles, bisex-

ual or unisexual (plant polygamo-monoecious).

Sepals four, decussate, imbricate; petals four,

free, imbricate Stamens eight, free, in two cycles.

Anthers deeply sagittate, dorsiﬁ xed, introrse, and

versatile, with a broad connective and a short

protrusion, opening longitudinally. Pollen grains

3-colporate, occasionally 4-ruporate. Nectary at

base of gynoecium well developed on sepaline

radii. Gynoecium of four carpels, they are con-

genitally united for almost their entire length,

free only at the very apex, with short style,

stigma large, capitate, the four free carpel tips

are postgenitally connected and form a single

stigmatic surface (Matthew and Endress 2005).

Ovary 4-locular and with axile placentation in

the lower part, but in the middle and upper parts

the partitions are incomplete and the ovules

borne on deeply intruded placentas. Ovules

numerous per locule, anatropous, Outer integ-

ument contributing to the micropyle zig-zag.

Fruits 4-locular, rather firm and dry berry.

Seeds arillate, with small, straight embryo, with

oily and proteinaceous endosperm and short,

ﬂ eshy funicle. Evergreen or deciduous shrubs,

small trees, or lianas; non laticiferous and with-

out coloured juice. The branchlets with large

pith. Cluster crystals and tanniniferous cells are

present in parenchymatous tissue. Nodes trila-

cunar with three traces. Leaves alternate, invo-

lute, simple, entire, glabrous or pubescent,

petiole bundle arcuate; stiuples free of one

another, caduceus, small, linear-lanceolate.

Stomata anomocytic. Ellagic acid, cyanidin, ﬂ a-

fonols (kaempferol and quercetin) present;

n = 12. . . . . . . . . . . . . . . . 1. stachyuraceae.

2 Inﬂ orescence cymose; sclereids absent; anthers

opening by apical pores or slits. Evergreen

trees; hairs unicellular. Leaves spiral or distic-

hous, estipulate, leaf margins crenate-serrulate.

Stomata anomocytic. Flowers in short dichasial

cymes, small, 5(6)-merous, sepals free, imbri-

cate; petals free or adnate to corolla, imbricate.

Stamens (8-)10(-15), in one cycle; ﬁ laments

broad, thick; anthers basiﬁ xed, setulose.

Carpels three; ovules two or many per locule.

Female gametophyte of Adoxa-type. Fruits dry,

ribbed, with papery exocarp and crustaceous

endocarp, eventually splitting septicidally into

three cocci. Seeds winged, with transparent

seed coat; embryo nearly straight, endosperm

copious; n = 24. . . . . . . . . . . 2. sladeniaceae.

3 Anthers basiﬁ xed; corc usually surﬁ cial; pseu-

dopollen absent; Fruits berries or drupes.

Evergreen trees or shrubs; hairs unicellular.

Leaves distichous or spiral, estipulate, leaf

margins entire, rarely serrate. Stomata anomo-

cytic. Flowers generally small, solitary, or in

clusters, bisexual or unisexual, 5-merous.

Sepals connate or distinct, calyx lobes always

persistent; petals distinct or connate basally,

thick or membranous. Stamens from 5 or 10

up to ca. 30, usually in one or two cycles; ﬁ la-

ments shorter or slightly longer than anthers;

anthers introrse, opening by longitudinal slits,

rarely by pore-like slits. Exine nearly smooth.

Carpels mostly (2)3–5(6); ovary superior to

inferior; ovules few or numerous per locule.

Female gametophyte of Polygonum-type.

Fruits berries or pomelike with succulent peri-

carp. Seeds usually numerous, often small, not

winged; seed coat usually cartilaginous, very

thin; endosperm present, often copious;

embryo slender, curved or hippocrepiform;

radicle always longer than the narrow cotyle-

dons, incumbent. Contain proanthocyanini-

dins, ﬂ avonols and ellagic acid, n = 12–13, 15,

18, 20–25. . . . . . . . . . 4. ternstroemiaceae.

3 Anthers versatile; cork usually deep-seated;

pseudopollen produced from connective; cap-

sule with persistent column. Trees or shrubs.

Leaves spiral or distichous, simple, usually

coriaceous; leaf margins serrate. Stomata

mostly paracytic (except in Franklinia,

Pyrenaria spp.). Flowers generally large and

showy, usually solitary, axillary, bisexual;

Sepals ﬁ ve or more, imbricate, calyx lobes

persistent or deciduous; petals ﬁ ve, rarely

numerous, free or connate basally; stamens

generally more than 40, in 2–6 cycles, free,

rarely connate; ﬁ laments several times as long

as anthers. Exine mostly ﬁ nely granulate or

ﬁ nely reticulate. Carpels mostly (3-)5(-10);

ovary superior. Female gametophyte of

Polygonum-, or rarely Allium-type. Fruits dry,