Takhtajan Armen. Flowering Plants

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182 Subclass V. DILLENIIDAE

1830). 28/470. Widespread in tropical regions, mainly

in rain forests.

2.1 CALOPHYLLOIDEAE (including

Kielmeyeroideae)

Leaves alternate, with pellucid dots, rarely canals.

Flowers bisexual, unisexual, or polygamous. Stamens

free or united at base or apex; anthers sometimes with

large, apical glands. Stylodia free or united. Fruits

drupes or capsules. Seeds small to large, exarillate.

Embryo large, with united large cotyledons. – calo-

phylleae: Neotatea, Marila, Mahurea, Clusiella,

Kielmeyera, Caraipa, Haploclathra, Poeciloneuron,

Mesua, Kayea, Mammea, Calophyllum, endodesmieae:

Endodesmia, Lebrunia.

2.2 CLUSIOIDEAE

Trees or shrubs, sometimes epiphytic. Leaves oppo-

site, usually with canals. Flowers unisexual, rarely

polygamous. Stamens free or in groups. Stylodia free

or united. Fruits capsules or drupes. Seeds arillate or

exarillate. Embryo with large hypocotyl and vestigial

or no cotyledons. – clusieae: Clusia (including

Quapoya, Havetia, Renggeria, Havetiopsis, Oedema-

topus, Pilosperma, Decaphalangium), Dystovomita,

Tovomita (including Tovomitidium), Tovomitopsis,

Chrysochlamys; garcinieae: Allanblackia, Garci nia

(including Ochrocarpus).

2.3 MORONOBEOIDEAE (CHRYSOPIOIDEAE)

Leaves opposite. Flowers bisexual. Stamens in ﬁ ve

groups or united into a tube. Style long, 5-lobed above.

Fruit mostly a berry. Seeds exarillate. Embryo undif-

ferentiated. – Pentadesma, Montrouziera, Moronobea,

Platonia, Symphonia, Thysanostemon.

2.4 LOROSTEMONOIDEAE

Flowers bisexual.– Lorostemon

Closely related to the Bonnetiaceae. In many respects,

on the basis of pollen morphology, Moronobeoideae dif-

fers markedly from the other subfamilies and in Erdtman’s

(1952) opinion possibly deserves a family rank.

3. HYPERICACEAE

A.L. de Jussieu 1789 (including Ascyraceae Plenck

1796). 9/550. Subcosmopolitan, mainly in temperate

regions and mountains of the tropics.

vismieae: Vismia, Harungana (including Psoro-

spermum); hypericeae: Hypericum, Lianthus,

Triadenum, Thornea, Santomasia; cratoxyleae:

Cratoxylum, Eliea.

The Hypericaceae differ from Clusiaceae in their

constantly bisexual ﬂ owers, and only very rarely do

the leaves have the numerous close, parallel nerves

or the wormlike secretory cells so characteristic of

Clusiaceae. These in Hypericaceae take the form of

translucent dots so familiar in the type genus Hypericum

and in this and other genera’s black dots. Besides, it is

not less important that in the Hypericaceae the stylodia

are more or less free. Both the Hypericaceae and

Clusiaceae differ markedly from the Bonnetiaceae in

the presence of a latex system.

4. ELATINACEAE

Dumortier 1829 (including Cryptaceae Raﬁ nesque

1820). 2/35. Nearly cosmopolitan. Bergia primarily in

tropical and subtropical regions and mainly of the Old

World, while Elatine occurs in all continents except

Antarctica.

Bergia, Elatine.

Elatinaceae have many features in common with

the Clusiaceae and especially with the Hypericaceae,

which also contain some aquatic forms (Cambessedes

1829; Takhtajan 1966; Corner 1976; Cronquist 1981;

Melikian 1992). The common features include oppo-

site leaves, syncarpous gynoecium with free stylodia

and capitate stigmas, septicidal capsules as well as

seed anatomy (Corner 1976; Melikian 1992) and stem

anatomy (Carlquist 1984). The reticulate sculpturing

of the seed coat in the Elatinaceae is very much like

that of some Clusiaceae and Bonnetiaceae (especially

the genus Ploiarium).

Bibliography

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Bonnetiaceae, Theaceae, and Guttiferae. In: P Baas, AJ

Bolton, and DM Catling, eds. Wood structure in biological

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Bennett GJ, H-H Lee, and TK Lowrey. 1989. Xanthones from

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Cambessedes J. 1829. Note sur les Elatinees: Nouvelle famille

des plantes. Paris.

Superorder ERICANAE 183

Carlquist S. 1984a. Wood and stem anatomy of Bergia suffruti-

cosa: relationships of Elatinaceae and broader signiﬁ cance

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seed anatomy, vol. 4, pp. 11. Nauka, St. Petersburg (in

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New York.

Order 53. OCHNALES

Trees or shrubs, sometimes lianas (some Quiinaceae),

less often subshrubs, very rarely herbs (some species of

Sauvagesia). Cristarque cells and mucilage cells and

canals usually present. Vascular bundles of the petiole

mostly cylindrical. Young stems of woody members

commonly with cortical vascular bundles and sometimes

with medullary bundles as well. Vessels mostly with

simple perforations, rarely with both simple and scalari-

form perforations, but Strasburgeriaceae have exclu-

sively scalariform perforations. Pits vestured. Rays

heterogeneous or homogeneous. Axial parenchyma usu-

ally apotracheal or metatracheal, often scanty. Nodes

mostly trilacunar or multilacunar. Leaves alternate or

less often opposite or verticillate (Quiinaceae), simple or

very rarely pinnately compound (Rhytidanthera splen-

dida), pinnately veined, often with numerous parallel

secondary veins, stipulate or rarely (Medusagynaceae)

estipulate. Stomata paracytic or less often anomocytic.

Flowers of medium size or small, in cymose or racemose

inﬂ orescences, bisexual, actinomorphic or rarely with a

184 Subclass V. DILLENIIDAE

tendency to zygomorphy (Luxemburgia). Perianth usu-

ally cyclic, rarely spirocyclic (Ochna and Ouratea of the

Ochnaceae and Strasburgeriaceae). Sepals (3)5(-12),

imbricate, free or more or less united. Petals (3)5(-12),

free, contorted or imbricate. Stamens few to many,

free, sometimes on elongated androphore; ﬁ laments free

or connate basally, ﬁ liform or broad and thick

(Strasburgeriaceae); anthers tetrasporangiate, basiﬁ xed

or dorsiﬁ xed, opening by longitudinal slits or apical

pores. Tapetum secretory. Microsporogenesis simulta-

neous. Pollen grains 2-celled or 3-celled, mostly

3-colporate or 3-porate. Sometimes the innermost sta-

mens sterile and transformed into a tube around the

ovary, or they form a lobed intrastaminal disc. Gynoecium

paracarpous; stylodia free or more often united into a

simple style that is sometimes lobed at the end; stigma

capitate; in each locule or on each parietal or axial

(Medusagynaceae) placenta 1(-2) to many ovules.

Ovules anatropous, campylotropous, or more or less

hemitropous, bitegmic or rarely (Lophiraceae) uniteg-

mic, tenuinucellate or rarely (Strasburgeriaceae) crassi-

nucellate. Female gametophyte of Polygonum- or

Allium-type. Endosperm nuclear. Fruits of different

types. Seeds with or without endosperm; embryo straight

or curved. Often producing proanthocyanins. Solitary or

clustered crystals of calcium oxalate frequently present

in some of the parenchymatous cells.

Related to the Hypericales.

Key to Families

1 Leaves usually alternate. Stylodia united into a

style. Cortical bundles present.

2 Stipules intrapetiolar, basally connate. Ovules

crassinucellate. Small to medium-sized trees.

Leaves alternate, simple, large, leathery, coria-

ceous, with very ﬁ ne, widely spaced teeth along

the margins of the distal half of the lamina; peti-

oles with narrow, lateral wings. Stomata anomo-

cytic. Mucilaginous cells containing a frothy,

red-staining contents present in both vegetative

and reproductive organs. Young stem with corti-

cal vascular bundles. Nodes predominantly trila-

cunar (nodes from older stems infrequently

pentalacunar) with three traces. Vessel elements

extremely long with exclusively scalariform per-

forations in almost vertical end walls and with

18–38 narrow and closely spaced bars; lateral

pitting opposite. Fibers (ﬁ ber- tracheids) extremely

long, with numerous bordered pits. Rays hetero-

geneous. Axial parenchyma apotracheal (diffuse

and scanty). Secondary phloem with primitive

sieve-elements and without ﬁ bers. Flowers

large, solitary, axillary, bisexual, actinomorphic.

Sepals 8–10, free, imbricate, spirally arranged,

gradually increasing in size from outer to inner,

persistent. Petals ﬁ ve, free, imbricate. Stamens

10, free, borne in a single cycle; ﬁ laments broad

and thick; anthers, dorsiﬁ xed, latrorse, opening

longitudinally. Pollen grains 3-colporate, distinctly

brevicolpate, psilate to psilate granular, tectate.

Disc intrastaminal, thick, annular at the base,

10-lobed above, lobes alternating with the sta-

mens. Gynoecium of 4–7 laterally completely

concrescent carpels that are also adnate with a

central column; style subulate, twisted, elongate,

terminates in lobed stigma; ovary angled,

4–7-locular, with one large ovule in each locule.

Ovules anatropous, epitropous ventral (ovules

pendulous, micropyle above, raphe ventral or

toward bundle), bitegmic, crassinucellate, with zig-

zag micropyle. Endosperm cellular. Fruits large,

woody, multiloculate, indehiscent, with persis-

tent sepals and style. Seeds somewhat ﬂ attened,

exotestal, with a straight embryo surrounded by

oily endosperm approximately ten cells in thick-

ness; sometimes with a rudimentary aril on funi-

cle; n = 250. . . . . . . . . .1. strasburgeriaceae.

2 Stipules extrapetiolar. Petals contorted or seldon

imbricate. Ovules tenuinucellate, bitegmic or rarely

(Lophiraceae) unitegmic. Vessels with simple or

occasionally with both simple and scalariform per-

forations. Lateral pitting alternate.

3 Style gynobasic, persistent even beyond ftuitfall.

Gynoecium is deeply lobed and the receptacle so

enlarged that the carpels appear to be free except

for their common style (Ochneae), or receptacle is

not distinctly enlarged and the carpels do not

appear distinct (Elvasieae). Fruits of separate

1-seeded drupes (Ochneae) or coriaceous and

deeply lobed, with one seed in each lobe (locule).

Seeds without endosperm, with highly vascu-

larized thin-walled, exotestal seed coat. Flowers

bisexual. Sepals ﬁ ve (Ochneae) or 3–6 (Elvasieae),

persistent. Petals ﬁ ve (Ochneae) or 3–6 (Elvasieae),

cadu cous. Stamens (8)10 to numerous, free, in

one or more cycles; anthers basiﬁ xed, more or

less latrorse, opening by longitudinal slits or

apical pores. Pollen grains 3-colporate. Leaves

Superorder ERICANAE 185

distichous, with short but distinct petioles. Stomata

paracytic. Vasicentric tracheids absent. Rays het-

erogeneous. Axial parenchyma apotracheal.

Crystarque cells commonly present. Trees or

shrubs; n = 12, 14, 24. . . . . . . . . . 4. ochna ceae.

3 Style apical. Carpels 2–5, with two to many

ovules per carpel. Ovules bitegmic or unitegmic.

Fruits berries or capsules. Receptacle not dis-

tinctly enlarged. Weeds with or without

endosperm. Flowers bisexual or functionally

polygamous. Petals ﬁ ve. Stamens ﬁ ve or numer-

ous, free or connate at the base. Leaves not

distichous. Vasicentric tracheids present or

absent. Crystarque cells usually absent Rays het-

erogeneous or homogeneous. Axial parenchyma

paratracheal of metatrachel.

4 Ovules bitegmic (outer integument of two lay-

ers, inner one of three layers), anatropous.

Shrubs, subshrubs or herbs. Rays heteroge-

neous. Axiall parenchyma paratracheal and

often scanty. Mucilage cells present.

Vasicentric tracheids absent. Leaves simple,

conduplicate-ﬂ at, rarely (Rhytidanthera) com-

pound; stipules persistent. Flowers bisexual,

actinomorphic; sepals free, imbricate, usually

persistent; petals contorted; anthers extrorse,

basiﬁ xed, 2-locular. Stamens ﬁ ve, free or con-

nate at the base, surrounded by numerous sta-

minodia (Sauvagesia). Carpels 2, 3, 5, with

1–2 (Euthemis) or many ovules per carpel.

Fruits berries (Euthemis) or capsular. Seeds

small, winged or not, with ﬁ brous exotegmen,

rarely (Indovethia) with long funiculus;

exotesta with large cells; endosperm ﬂ eshy.

n = 18.. . . . . . . . . . . . . . .2. sauvagesiaceae.

4 Ovules unitegmic (with two completely fused

integuments), numerous (10–20) but only

one fertile, anatropous. Carpels two, with

5–10 ovules per carpel. Fruits 1-seeded cap-

sules, with two persistent outer sepals

enlarged and winglike. Seeds large, without

endosperm, with testa reduced to approxi-

mately three layers of unspecialized cells,

compressed between pericarp and large

embryo. Stamens numerous, in 3–5 cycles.

Axial parenchyma metatracheal. Rays homo-

geneous. Mucilage cells absent. Vasicentric

tracheids present. Stomata anomocytic. Tall

trees; n = 14. . . . . . . . . . . . 3. lophiraceae.

1 Leaves opposite or verticillate. Cortical bundles

absent.

5 Petals contorted. Small, glabrous trees up to 10 m

with a rounded crown. Vessels with simple perfo-

rations in slightly inclined to nearly vertical end

wall; lateral pitting from opposite to predomi-

nately alternate. Fibers with small bordered pits.

Rays heterogeneous. Axial parenchyma diffuse

and diffuse-in-aggregates. Secondary phloem

stratiﬁ ed into alternating concentric rings of sieve

tissue and sclerenchyma. Nodes pentalacunar with

ﬁ ve traces. Leaves opposite, leathery, and shiny,

simple, often emarginate, margins crenate with

ﬁ ne, widely spaced teeth, pinnately veined, with a

pair of colleters in the leaf margins and in the leaf

axils external to the axillary bud, with scattered

mucilaginous cells in the mesophyll; stipules

wanting, petiole bundles arcuate. Stomata anomo-

cytic. Flowers medium-sized in lax, terminal,

cymose inﬂ orescences, andromonoecious (bisex-

ual and exclusively male usually within the same

inﬂ orescence), actinomorphic with an elongate

receptacle foetid. Sepals ﬁ ve, connate at the base,

imbricate, reﬂ exed persistent. Petals ﬁ ve, free,

basally thickened, spatulate, contorted (with each

petal having one margin exterior and the other

margin interior), at anthesis reﬂ exed. Stamens

very numerous, free, spirally arranged on an elon-

gated receptacle, and characterized by ﬁ ve major

stamen fascicle traces (stamen trunk bundles); ﬁ l-

aments very slender, shorter than the petals.

Anthers basiﬁ xed, usually with a small, apical

extension of the connective, 2-locular, the locules

often set at different heights, opening longitudi-

nally. Pollen grains 3-porate, tectate-columellate,

with short and rather wide columellae and the

massive foot layer, forming about one half of total

exine thickness; the apertures protrude and are

formed by a tapering, conelike extension of the

exine; ornamentation consists of very ﬁ ne striae.

Gynoecium of 16–19 (up to 25) carpels fused ven-

trally to a solid parenchymatous central column (a

prolonged receptacle). Below the level of ovular

insertion the ventral sutures are open although car-

pels are always fused to the central column.

Stylodia rather short, forming a ring around the

shoulders of the ovary persistent; stigmas capitate,

disclike, with a papillate upper surface; ovary with

two ovules in each locule on axile placentas, upper

186 Subclass V. DILLENIIDAE

one ascending or erect (micropyle below), the

lower one descending or hanging (micropyle

above). Ovules large, with long and curved funiculi,

anatropous, bitegmic, with a straight micropyle,

vascularized by a single bundle that extends

unbranched from the funiculus to the chalaza, with

a very thin nucellus composed of one cell layer at

maturity. Fruit a slightly woody, septicidal pendu-

lous capsule; each carpel separates acropetally

from the central column, with only a distal connec-

tion maintained; the carpels thus diverging from

the top of the central column like the ribs of an

umbrella; dehiscence of the individual carpels is

along the entire ventral suture. Seeds two in each

locule, small, compressed, winged all around,

exarillate, with straight embryo and narrow layer

of endosperm consisting of a cellular zone of very

thin-walled cells without endosperm. The mature

seed coat thin, consisting of four or ﬁ ve layers of

cells and exotegmic with ﬁ bers. Tanniniferous.

Dark-staining phenolic compounds throughout the

plant body. . . . . . . . . . . . . . 5. medusa gyn a ceae.

5 Petals imbricate. Evergreen trees, sometimes to

30 m tall, shrubs, or sometimes woody lianas.

With scattered secretory cells in the parenchyma-

tous tissues of the young stem and with lysigenous

mucilage-channels, especially within the vascu-

lar strand of the midrib and petioles of the leaves.

Cortical bundles absent. Nodes trilacunar.

Vessels with simple or seldom scalariform

perforations. Fibers with evidently bordered

pits. Rays heterogeneous. Axial parenchyma

apotracheal diffuse or sometimes banded. Leaves

simple or (Touroulla and Froesia) pinnately com-

pound; secondary veins numerous, tertiaries

feather-veined; veins, especially the smaller

ones, sheathed by very thick-walled ﬁ bers;

stipules interpetiolar, usually conspicuous, folia-

ceous or setose, entire or divided into several

setose parts, persistent, pubescent. Stomata para-

cytic. Flowers in axillary or terminal panicles or

racemes, bisexual or unisexual (plants androdi-

oecious in Quiina and Touroulia or dioecious in

Lacunaria), actinomorphic. Sepals 4-4 small,

unequal, imbricate. Petals 4–5(-8), imbricate.

Stamens 9 to ca. 100 in Lacunaria, Quiina, or

sometimes over 300 in Froesia (Zizka et

Schneider 2004); ﬁ laments ﬁ liform, free or con-

nate at the base, and sometimes adnate to the

base of the petals; anthers small, basiﬁ xed,

2-locular, opening longitudinally. Pollen grains

3-colporate. Gynoecium of three free carpels

(Froesia) or more often of 2–14 united carpels

with 2-14-locular ovary and linear stylodia; stig-

mas obliquely peltate. Ovules paired, ascending,

anatropous. Fruits berrylike or 1–3 follicles in

Froesia. Seeds one to more than 20 in berrylike

fruits of one per follicle, hairy, or glabrous in

Froesia, without endosperm and with straight

embryo. . . . . . . . . . . . . . . . . . . . .6. quiinaceae.

1. STRASBURGERIACEAE

Solereder 1908. 1/1. New Caledonia.

Strasburgeria.

Strasburgeria robusta is one of the most remarkable

New-Caledonian relicts still preserving primitive char-

acters, such as very long vessel elements with 20–30

bars, spirally arranged sepals, crassinucellate ovules,

etc. But like many other phylogenetic relicts, it is very

heterobathmic and has a number of specialized

characters, especially in pollen, gynoecium, and fruit

morphology. It is an early side branch from the

ochnalean ancestral stock.

2. SAUVAGESIACEAE

Dumortier 1829 (including Euthemidaceae Sole-

reder 1908, Luxemburgiaceae Solereder 1908).

20/120. Pantropical, but mainly represented in the

Neotropics.

2.1 LUXEMBURGIOIDEAE

Tree. Flowers yellow; sepals caducous; androecium

obliquely zygomorphic in bud, only adaxial; ﬁ laments

more or less connate; pollen exine with small perfora-

tions. Gynoecium of three carpels. – Luxemburgia.

2.2 SAUVAGESIOIDEAE

Trees, shrubs, subshrubs, or herbs. Rays heteroge-

neous. Axial parenchyma paratracheal and often

scanty. Secondary veins archingly joined near the leaf

margin. Staminodia 10-numerous in one or more

cycles. Stamens five, more or less adnate at the

base to a short tube of inner staminodia. Anthers

opening by two lateral slits. Gynoecium of 2–3 carpels.

Superorder ERICANAE 187

Ovary 1-locular with intruding placentas and

numerous ovules. Fruits a coriaceous capsule. Seeds

usually numerous, small. – Godoya, Rhytidanthera,

Krukoviella, Cespedesia, Testulea, Fleurydora,

Philacra, Blastemanthus, Poecilandra, Wallacea,

Schuurmansia, Schuurmansiella, Tyleria (including

Adenanthe?), Adenanthe, Adenarake, Indosinia, Sinia,

Sauvagesia (including Indovethia?).

2.3 EUTHEMIDOIDEAE

Shrubs. Numerous parallel secondary veins reaching

denticulate leaf margin. Staminodia 0(-5), ﬁ lamentous.

Stamens free. Anthers opening by an apical pore.

Gynoecium of ﬁ ve carpels. Ovary 5-locular, with 1–2

ovules per locule. Fruits berries with ﬁ ve, usually 1-seeded

pyrenes. Seeds one or two per locule. – Euthemis.

Sauvagesiaceae are usually included in the

Ochnaceae, which makes the later very heterogeneous.

They have no close afﬁ nity with the Ochnaceae and

differ markedly from them in many respects, including

seed coat anatomy (Corner 1976; Takhtajan 1987).

3. LOPHIRACEAE

Loudon 1830. 1/2. Tropical Africa.

Lophira.

A specialized unigeneric family that differs mark-

edly from both the Sauvagesiaceae and Ochnaceae, but

it is nearer to the Sauvagesiaceae.

4. OCHNACEAE

A.P. de Candolle 1811 (including Gomphiaceae A.P.

de Candolle ex Schnizlein 1843–1870). 6/480. Pantro-

pical with only a few representatives in the subtropics

(some species of Ochna partly cross the Tropic of

Cancer in northern India, to ca. 30° N). The tribe

Elvasieae is Neotropical.

ochneae: Ochna, Brackenridgea, Ouratea (including

Idertia), Gomphia (including Rhabdophyllum?, Campylo-

spermum?), Perissocarpa; elvasieae: Elvasia.

5. MEDUSAGYNACEAE

Engler et Gilg 1924. 1/1. Seychelles Islands. Medusagyne.

Related to the Ochnaceae.

6. QUIINACEAE

Engler 1888. 4/50. Tropical America from Belize south

to Bolivia and southern Brazil, but mainly Amazon

baSinica.

Froesia, Quiina, Touroulia, Lacunaria.

Evidently close to the Ochnaceae (see Gottwald

and Parameswaran 1967).

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Order 54. ERICALES

Shrubs or less often small trees, perennial herbs or

woody lianas. Raphide mostly cells absent. Vessels

mostly with scalariform perforations, sometimes with

many bars (up to 50 in Clethraceae and up to 65–70 in

Cyrillaceae), but often with both scalariform and sim-

ple perforations or only with simple perforations;

lateral pitting from scalariform to alternate. Fibers

with bordered pits. Rays typically heterogeneous.

Axial parenchyma diffuse, often scanty or wanting.

Sieve-element plastids of S- or (Cyrillaceae) Pcf-type.

Nodes unilacunar, trilacunar (some Ericaceae and

some Styphelioideae) or multilacunar (some Styphe-

lioideae). Leaves alternate, less often opposite or

verticillate, simple, entire, estipulate, sometimes

without chlorophyll and reduced to mere scales.

Stomata anomocytic, paracytic, tetracytic, or of other

types. Flowers in various (mostly racemose) inﬂ ores-

cences or sometimes solitary (axillary or terminal),

bisexual or rarely unisexual, usually actinomorphic.

Sepals (3-)5(-7), free or basally connate, imbricate or

valvate. Petals isomerous with sepals, free or more

often connate into sympetalous corolla with usually

imbricate or contorted, more rarely valvate lobes;

rarely petals wanting. Stamens mostly twice as many

as the petals or less often the same number and

then alternate, rarely only two (Ceratiola in the

Empetroideae) or up to 20; ﬁ laments free or more or

less adnate to the corolla; anthers tetrasporangiate or

disporangiate, usually inverted, opening longitudinally

or much more often by apical (morphologically basal)

pores or slits, often with two or more slender append-

ages, which are rarely borne on the upper part of the

ﬁ lament instead of on the anther. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains 2-celled

or sometimes 3-celled, very often in tetrads, mostly

3-colporate. Nectary disc usually present, intrastaminal.

Gynoecium of 2–10 united carpels, with usually

simple style; ovary superior or inferior, 1–10-locular,

with numerous or rarely several or even one ovule

per carpel. Ovules anatropous or hemitropous, uniteg-

mic, tenuinucellate. Female gametophyte usually of

Polygonum-type. Endothelium present. Endosperm

usually cellular (nuclear in Rhododendron and Vacci-

nium), with terminal haustoria that are usually micro-

pylar and chalazal. Fruits septicidal or loculicidal

capsules, berries, or drupes. Seeds mostly small or

Superorder ERICANAE 189

minute, exotestal, with copious or scanty endosperm

and small or minute embryo. Often producing iridoid

compounds, alkaloids present (rarely), or absent.

Key to Families

1 Raphide cells present. Trees, shrubs, or woody

lianas; hairs often more or less ﬂ attened setose. Pith

often septate. Vessel elements with spiral thicken-

ing, their ends more or less oblique, with scalari-

form perforations that have 15 and more bars or with

mixed scalariform and simple perforations; lateral

pitting opposite or less often scalariform or alter-

nate. Fibers mostly long, with distinctly or dared

pits. Nodes unilacunar or seldom trilacunar. Leaves

dentate or entire. Stomata anomocytic or sometimes

nearly paracytic. Flowers small, in axillary or lat-

eral, basically cymose inﬂ orescences (which some-

times reduced to one or a few ﬂ owers), often on old

wood, bisexual, polygamous or dioecious, often

subtended by small bracteoles. Sepals and petals

mostly ﬁ ve each, sometimes four, seldom up to

seven, usually quincuncial in bud, free or connate at

the base. Stamens branched, or unbranched; matur-

ing centrifugally; free of the perianth; all equal; free

of one another, or coherent (then variously united at

the base, often in ﬁ ve clusters opposite the petals);

numerous to 10, often in ﬁ ve antepetalous fascicles,

that may adnate to the base of the petals. Anthers

erect or inﬂ exed in bud, versatile, inverted, opening

by morphologically basal but seemingly apical short

slits or pores. Pollen grains 2-celled, in monads or

sometimes (Saurauia elegans) in tetrads, with com-

plete tectum and reduced columellae, psilate or

granulate. Gynoecium of 3–5 or many (up to 30 or

more) united carpels, with free or more or less con-

nate stylodia; ovary superior, completely or incom-

pletely trilacunar to multilacunar, with numerous or

(Clematoclethra) only 10 ovules in each locule on

axile or nearly axile placentas. Ovules anatropous,

hypostatic (with the distinctive thick-walled nucellar

area in the chalaza), with conspicuous endothelium

and a well-developed hypostase. Fruits berries, hairy

or glabrous, with massive placentae, or sometimes

more or less loculicidal capsules. Seeds small, aril-

late or not, with copious endosperm and well-

developed, straight or slightly curved embryo.

Iridoid compounds found in Actinidia; myricetin

was detected in leaves, actinidin, a proteinase simi-

lar to papain was detected in kiwifruits (McDowall

1970; Dressler and Bayer 2004). n = 30 (Saurauia,

South American species – Soejarto 1969, 1970) and

n = 20 (Asian species), 29 (Actinidia – Yan et al.

1997). . . . . . . . . . . . . . . . . . . . . 1. actinidia ceae.

1 Raphide cells absent.

2 Petals, when present, free or nearly so. Pollen

grains usually in monads. Leaves alternate.

Nodes only unilacunar.

3 Anthers deeply inverted, opening by slitlike

pores at the apparent apex. Intrastaminal disc

wanting, but the basal part of the ovary often

nectariferous. Sepals imbricate, persistent.

Petals ﬁ ve, free, imbricate. Stamens 10(12),

free. Anthers not appendages. Pollen grains

single or rarely in tetrads, 3-colporate. Gynoe-

cium of three carpels. Ovary superior, 3-locular,

ovules numerous on intrusive placentas, anatro-

pous. Fruits loculicidal capsules. Seeds numer-

ous, often winged. Seed coat present, exotestal,

consisting of a single layer of cells; embryo

straight, short, cylindrical; endosperm moder-

ate, ﬂ eshy Small trees or shrubs; indumentum of

simple and fasciculate, often stellate hairs.

Vessels with scalariform perforation with 20–50

bars, pits vestured. Alkaloids and iridoids not

detected. Kestose and isokestose oligosaccha-

rides present, n = 8. . . . . . . . . 2. clethraceae.

3 Anthers not obviously inverted, opening by

longitudinal slits or apical pores. Intrastaminal

nectary disc present. Sepals 5(6), free or rarely

very shortly connate, imbricate or rarely val-

vate, persistent. Petals 5(6), free or slightly

connected below, imbricate or contorted.

Stamens 5 or 10 in two whorls, ﬁ laments elon-

gate, sometimes broadened, free or adnate to

the corolla at the very base (Purdiaea). Pollen

grains 3(-4)-colporate. Gynoecium of 2–5 car-

pels. Ovary 2-4(5)-locular; ovules 1–3 per

locule, pendulous, apotropous. Funiculus

long. Fruits small, indehiscent, the pericarp

thin and dry, sometimes a samaroid nutlet.

Seeds usually without seed coat; embryo

straight, usually cylindrical, with small,

slightly expanded cotyledons; endosperm

copious, ﬂ eshy-hard. Small glabrous trees or

shrubs. Vessels with scalariform perforations

with 20-50(-70) bars. Leaves short-petioled or

sessile. Present myricetin, ellagic acid and pro-

cyanidin, n = 10. . . . . . . . . . . 3. cyrillaceae.

190 Subclass V. DILLENIIDAE

2 Petals, when present, mostly united into a sym-

petalous corolla. Pollen grains mostly in tetrads.

Mainly shrubs, less often perennial herbs, small

trees, or rarely trailing or scrambling lianas.

Indumentum unicellular and multicellular, or

none. Vessels mostly with scalariform perfora-

tion with many bars. Nodes multilacunar, trila-

cunar, or unilacunar. Leaves mostly with palmate

or subparallel venation, rarely with pinnate vena-

tion (Prionotes and Lebetanthus), alternate or

opposite. Stomata mostly anomocytic, but in

Vaccinioideae mostly paracytic. Flowers mostly

bisexual or rarely (Empetroideae) unisexual.

Sepals (3)4–7(8), mostly ﬁ ve, usually more or

less connate. Petals the same number as sepals,

more or less connate to form a short or conspicu-

ous corolla tube. Stamens mostly in two cycles,

twice as many as petals, seldom more numerous

(up to 20) or less often the same number as

petals. Anthers with exothecium or rarely

(Enkianthus) with endothecium, tetrasporangiate

and 2-locular (rarely 4-locular) or disporangiate

and usually 1-locular (Styphelioideae), opening

by pores at the apparent apex, or by elongate

slits, sometimes by longitudinally, often with

appendages. Pollen grains in tetrahedral tetrads

or rarely (Enkianthus) in monads. Intrastaminal

nectary disc present. Gynoecium of 2–10, mostly

ﬁ ve or, less often, four carpels. Ovary usually

superior or inferior, usually with the same num-

ber of locules as carpels and rarely (as in

Scyphogyne and related genera) unilocular, with

a solitary pendulous ovule; ovules 1-several or

more often numerous on each placenta, anatro-

pous to subcampylotropous. Fruits septicidal or

loculicidal capsules, or berries or drupes, seldom

a small nut. Seeds numerous, small to minute,

sometimes winged, embryo straight, fusiform,

rarely embryo minute, undifferentiated;

endosperm ﬂ eshy, with haustoria at both ends.

Present ﬂ avonoids, ﬂ avones, simple phenols,

anthocyanins, ellagic acid, and methyl salicylate;

iridoids detected.. . . . . . . . . . . . . . 4. ericaceae.

1. ACTINIDIACEAE

Gilg et Werdermann 1925 (including Saurauiaceae

Grisebach 1854). 3/365. Tropical and subtropical

regions of eastern and Southeast Asia, West Malesia,

America, and Australia (only one species of Saurauia

in northeastern Queensland); Actinidia (55) is distrib-

uted in the Himalayas and eastern and Southeast Asia,

Clematoclethra (1) in temperate China.

1.1 SAURAUIOIDEAE

Flowers usually bisexual, rarely functionally dioecious,

predominantly 5-merous. Petals usually connate at the

base. Stamens 13–50 or more, basally connate, and dis-

tinctly epipetalous to only partially so. Anthers opening

by rimiform pores. Gynoecium of 3–5 (very rarely seven)

carpels. Stylodia emerging from a depression at the apex,

free or united for three fourths of their length. Ovules

variable in number, commonly numerous in each locule.

Fruits usually typical berries with numerous small seeds

embedded in a mucilaginous pulp, n = 13. – Saurauia.

1.2 ACTINIDIOIDEAE

Flowers dioecious or polygamous, with 5-merous or

4-merous perianth. Petals free to basally connate. Stamens

numerous, free. Anthers opening by short subapical lon-

gitudinal slits or by rimiform pores. Gynoecium of 3–5 or

15 to slightly more than 30 carpels. Stylodia free. Ovules

numerous in each locule. Fruits berries, include about 40

locules and 1,500 seeds, n = 29. – Actinidia.

1.3 CLEMATOCLETHROIDEAE

Flowers dioecious or polygamous, basally 5-merous.

Stamens 10–30, free. Anthers opening longitudinally.

Gynoecium mostly of ﬁ ve or four carpels. Stylodia

united into a style arising from a conspicuous depres-

sion and terminated by a slightly swollen stigmatic

surface. Ovules about 10 per locule. Fruits capsules or

drupaceous, n = 12. – Clematoclethra.

I agree with Dickison (1972: 53) that “the similari-

ties between the Actinidiaceae and Ericales are impres-

sive and indicate that the close afﬁ nities of this family

may very well lie within the ericalean complex.”

Although in many respects, particularly in embryology

(Crété 1944), Actinidiaceae are closely related to other

families of Ericales (especially Clethraceae), they differ

in the presence of the raphides, stamen fascicle traces,

hypostatic ovules, the absence of terminal haustoria,

and some other less important features.

2. CLETHRACEAE

Klotzsch 1851. 1/85. Eastern and Southeast Asia,

Malesia, southeastern United States, central and tropical

Superorder ERICANAE 191

South America, with one endemic species on the island

of Madeira.

Clethra.

Clethraceae are related to the Actinidiaceae, which is

conﬁ rmed also by embryological data (Veilllet- Bar-

toszewska 1960). They reveal also deﬁ nite closeness to

the Theaceae, especially in the anatomy of the seed coat

and in ﬂ ower morphology, including the presence of

nectariferous tissues in the basal part of the ovary of

some species of Clethra, which resemble Eurya japon-

ica (Brown 1938). Thus, specialized nectariferous tis-

sues have already developed in Clethra, which gave rise

to the intrastaminal nectary disc of other Ericales

(Takhtajan 1966). Although Clethraceae have many fea-

tures in common with the Theales, especially with the

genus Eurya in the Theaceae, they are closer to the

Ericales, which is conﬁ rmed not only by the ﬂ ower mor-

phology but also by their unitegmic ovules, presence of

endothelium and endospermal haustoria (Poddubnaja-

Arnoldi 1982), anatomy of the seed coat (Corner 1976)

and wood characters (Giebel and Dickison 1976).

3. CYRILLACEAE

Endlicher 1841. 3/14. Central America, northern parts

of South America, West Indies, coastal plains of south-

eastern United States.

Cyrilla, Cliftonia, Purdiaea.

Jussieu (1789) placed Cyrilla (the only known cyril-

laceous genus at that time) in his Ericeae, which has

been followed by some other botanists including

Agardh (1858). According to Hallier (1912), Cyrrilla-

ceae originated from the Ericaceae and are so close to

them that they could even be considered a tribe of the

Ericaceae. This idea of the closeness of the Cyrillaceae

to the Ericaceae has been accepted by an increasing

number of botanists (including Wettstein [1935] and

Copeland [1953]) and by some later authors. Dahlgren

(1975, 1989), Thorne (1976, 1992a, b, 2000, 2006),

and Cronquist (1981, 1988) place the Cyrillaceae close

to the Clethraceae. These two families have many

similarities in gross morphology and in embryology,

palynology, and chemistry (including the absence of

iridoid compounds). Cyrilla and Clethra are also very

similar in aspect (Cronquist 1981). Anderberg (1993)

notes that the genus Purdiaea includes one taxon that

has formerly been placed in the Clethraceae, thus

reﬂ ecting the symplesiomorphic similarity of the two

families.

4. ERICACEAE

A.L. de Jussieu 1789 (including Andromedaceae

A.P. de Candolle ex Schnizlein 1843–1870, Arbutaceae

Bromhead 1840, Arctostaphylaceae J.G. Agardh

1858, Azaleaceae Vest 1818, Empetraceae Berchtold

et J. Presl 1820, Epacridaceae R. Brown 1810,

Hypopityaceae Klotzsch 1851, Ledaceae Gmelin

1803, Menzieziaceae Klotzsch 1851, Monotropaceae

Nuttall 1818, Oxycoccaceae A. Kerner 1891, Priono-

taceae Hutchinson 1969, Pyrolaceae Lindley 1829,

Rhododendraceae A.L. de Jussieu 1789, Rhodoraceae

Ventenat 1799, Salaxidaceae J.G. Agardh 1858,

Styphe-liaceae Horaninow 1834, Vacciniaceae

A.P. de Candolle ex Perleb 1818). 128/c.4000. Widely

distributed in almost all parts of the world with large

concentrations in the Himalayas and southwestern

China, New Guinea and southern Africa; absent from

most of Australia.

4.1 ENKIANTHOIDEAE

Usually deciduous or sometimes evergreen shrub. Pith

heterogeneous (pith with small, thick-walled and ligni-

ﬁ ed and larger and thin walled cells mixed). Leaves

ﬂ at and fairly large, serrulate or entire. Flowers in

raceme- or umbel-like inﬂ orescences. Corolla urceo-

late to companulate. Stamens with papillose to hairy

ﬁ laments, anthers with paired awns (ﬁ brous endothe-

cium), which gradually narrow from the thecae. Pollen

grains trinucleate, in monads, without viscin threads.

Ovary superior, 5-locular. Fruits loculicidal capsules,

opening with ﬁ ve valves. Seeds rather large, often

winged on the margins. n = 11. – Enkianthus.

4.2 MONOTROPOIDEAE

Strongly mycotrophic and more or less fleshy

perennial herbs devoid of chlorophyll. Multicellular

hairs present. Leaves reduced to scales. Corolla

persistent, rarely absent. Anthers sometimes spurred at

the morphological base, without viscin threads, open-

ing by longitudinal slits or sometimes by terminal

pores or by slits across the broad locular summits.

Pollen grains in monads. Ovary 1–6-locular. Fruits

loculicidal or septicidal capsules, or baccate and inde-

hiscent; embryo minute, undifferentiated, n = 8, 13,

19, 23, etc. – pterosporeae: Pterospora, Sarcodes,

Allotropa; pleuricosporeae: Pleuricospora; mono-

tropeae: Cheilotheca, Monotro psis, Monotropa,

Monotro pastrum; hemitomeae: Hemitomes, Pityopus,

Hypopitys.