Takhtajan Armen. Flowering Plants

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222 Subclass V. DILLENIIDAE

anthers 2-locular, opening longitudi-

nally. Staminodes 5–7, in the same

series as the fertile stamens, or inter-

nal to the fertile stamens. Pollen

grains 3-colporate. Gynoecium of

three carpels with simple style ter-

minated by small capitate stigma.

Ovary superior, 1-locular above, per-

fectly 3-locular at the base, with two,

ascending ovules per carpel but only

one developing. Fruits tardily dehis-

cent 1-locular, tomentose, 1-seeded

capsules. Seed small, oblong with a

persistent style, surrounded by the

persistent calyx, corolla and androe-

cium; seed coat with palisade tissue

underneath collapsed polygonal

cells. . . . . . 6. dipentodonta ceae.

8 Placentation parietal.

9 Petals 5.

10 Anthers basiﬁ xed. Shrubs or

herbs (mostly perennial), less

often small trees or scandent.

Vessels with scalariform or (espe-

cially in herbs) simple per-

forations; lateral pitting from

opposite to scalariform, some-

times alternate. Fibers with sim-

ple or bordered pits, septate.

Axial parenchyma absent or

very scanty. Leaves alternate or

sometimes opposite (some spe-

cies of Hybanthus and Rinorea)

simple, entire or toothed, rarely

lobed, with leafy or small stipules.

Stomata paracytic or anisocytic.

Inﬂ orescences axillary racemes,

spikes, panicles or the ﬂ owers

solitary. Flowers actinomorphic

or more or less zygomorphic,

bisexual or rarely polygamous or

dioecious. Sepals ﬁ ve, often per-

sistent, imbricate, or open in bud.

Petals ﬁ ve, imbricate or con-

torted, clawed or sessile, in zygo-

morphic ﬂ owers unequal, the

lowest one usually prolonged

behind into a spur. Stamens ﬁ ve

(three in Leonia triandra), alter-

nating with the corolla; ﬁ laments

very short, free or more or less

connate; anthers usually more or

less connivent in a ring around

the ovary, introrse, with a con-

nective often prolonged into a

membranous appendage, open-

ing longitudinally. Pollen grains

3(-5)-colporate. Gynoecium of

(2)3–5 carpels, with stylodia

united into a style; the stigma

simple or lobed. Ovary superior,

unilocular, with 1–2 or many,

anatropous ovules on (2)3–5

parietal placentas. Fruits loculi-

cidal and valvular capsules or

sometimes berries, rarely

(Leonia) nutlike. Seeds winged

or wingless, with straight, green

(some species of Viola) embryo

and ﬂ at cotyledons surrounded

by copious, oily, rarely scanty

endosperm, often arillate. Contain

ﬂ avonols (kaempferol and quer-

cetin), alkaloids present or absent,

inulin recorder in Ionidium; n =

6–13,17, 21, 23.. . 5. viola ceae.

10 Anthers dorsiﬁ xed. Herbs,

shrubs, or rarely trees, erect or

decumbent, usually with glandu-

lar or non-glandular trichomes.

Vessels with simple or less often

with simple or scalariform perfo-

rations; lateral pitting alternate.

Rays heterogeneous or homoge-

neous. Axial parenchyma apotra-

cheal. Leaves alternate, simple,

entire or lobed, often with two

glands or extraﬂ oral nectaries at

the back of the lamina; stipules

small (Erblichia), or lacking

(usually, but often with a pair of

glands or extraﬂ oral nectaries

at the base of the blade). Stomata

of various types. Inﬂ orescences

axillary racemes or cymes or

ﬂ owers solitary. Flowers bisexual,

actinomorphic, with a short to

more often tubular hypanthium,

Superorder VIOLANAE 223

often subtended by two bracte-

oles. Calyx tubular, 5-toothed,

teeth imbricate. Petals ﬁ ve,

inserted on the calyx tube, free,

clawed, contorted in bud, usually

ephemeral. Hypanthium some-

times with fringed corona or ﬁ ve

glands or lobes between corona

and androecium. Stamens ﬁ ve,

alternating with the corolla, free

or partially attached in throat of

hypanthium. Pollen grains in

monads, usually 3-colporate.

Gynoecium of three carpels; sty-

lodia free, opposite the placentas,

sometimes biﬁ d, often with a

fringed or brushlike commissural

stigma; ovary superior or some-

times semi-inferior, 1-locular;

ovules numerous on each parietal

placenta, anatropous. Fruits cap-

sules, opening loculicidally into

three valves with the placenta in

the middle of each. Seeds pitted,

arillate, endotestal cells large,

exotegmen palisade; embryo

large, straight or weakly curved,

endosperm copious, oily; coty-

ledons two, planoconvex.

Producing cyclopentenoid cyano-

hydrins glycosides, linamarin,

and sometimes alkaloids, n = 5,

7, 8(10), 13. . . 9. turneraceae.

9 Petals six. Small dioecious trees with

soft wood. Vessels with very oblique

end walls but usually with simple

perforations. Lateral pitting alter-

nate. Fibers with strongly thickened

walls and with large bordered pits

and numerous thin cross-partitions.

Rays heterogeneous. Axial paren-

chyma nearly or quite wanting.

Nodes trilacunar. Leaves alternate,

simple, distichous, serrate or dentate,

with very small, caducous stipules.

Stomata paracytic. Inﬂ orescences

terminal and axillary racemes or

panicles. Flowers actinomorphic,

3-merous. Perianth in male ﬂ owers

tubular, 6-lobed, the lobes in two

series, imbricate (the outer series

probably represents a calyx, and the

inner one a corolla (Swamy 1953).

Three large, ﬂ eshy nectary glands

present between the perianth and the

androecium. Stamens united into a

column with clavate common pro-

longed apical connective. Anthers

2-locular, with a prolonged connec-

tive, extrorse, opening longitudi-

nally. Pollen grains 3-colpate.

Perianth of the female ﬂ owers of

three free sepals alternating with

three free petals. Gynoecium of 8–12

carpels. Stigma sessile, thick, dis-

coid, with as many radiating ridges

as carpels and with the aperture in

the middle. Ovary superior, 1-locu-

lar, more or less septate near the

summit but unilocular below, with

numerous (30–100), basal, erect

ovules on prominent funicles. Fruits

ﬂ eshy loculicidal capsules, or ulti-

mately dehiscing into 8–12 reﬂ exed

segments, enclosing numerous

seeds. Seeds arillate, perisperm

present in a very thin layer, embryo

fairly large, straight; endosperm

rather scanty, oily, cotyledons 2,

n = 9.. . . . . . 7. scyphostegia ceae.

7 Stamens usually more than ﬁ ve. Trees

and shrubs, rarely with stellate indu-

mentum (Kiggelaria, Rayania). Vessels

with scalariform or more often simple

perforations. Fibers with simple or bor-

dered pits, usually septate. Axial paren-

chyma absent. Leaves alternate or

rarely opposite (Pseudoscolopia,

Abatia, Apherema), simple, often dis-

tichous and coriaceous, serrate or entire,

sometimes pellucid-dotted or lined,

stipules usually present, often cadu-

cous, rarely absent (Abatia, Bembicia).

Stomata paracytic or anisocytic.

Inﬂ orescences axillary, terminal or

below the leaves, racemes, cymes,

rarely (Bembicia) in short axillary

conelike heads surrounded by closely

224 Subclass V. DILLENIIDAE

imbricate leathery bracts, the outer ones

empty; bracteoles 2-keeled, sometimes

adnate to the midrib with the ﬂ owers

epiphyllous (in Phyllobotryon,

Phylloclinium, Moc querysia). Flowers

bisexual or unisexual. Perianth parts

clearly differentiated into sepals and

petals or rarely (Erythrospermeae)

poorly differentiated. Sepals imbricate

or rarely valvate, persistent. Petals

present or absent (Flacourtieae,

Casearieae, Bembicieae). Petals with

an adaxial scale (in some genera of

Erythrospermeae and in Pangieae).

Stamens development centrifugally or

centripetally, numerous or occasionally

equal in number to the petals, the ﬁ la-

ments distinct or grouped into fascicles

opposite the petals. Anthers linear to

about as long as wide, dorsiﬁ xed or

basiﬁ xed, usually latrorse, sometimes

the connective produced apically, open-

ing usually longitudinally, or rarely

(Kigellaria) via pores. Pollen grains

3(-6)-colporate. Prismatic crystals in

ray cells usually present (except for

Poliothyrsis). Hypogenous disc

extrastaminal or intrastaminal, or sepa-

rate members or annular (Bembicia).

Gynoecium of 2–10 carpels. Ovary

superior or rarely (Bembicia) inferior,

with 2(3)–5(-10) placentas (sometimes

appearing almost basal) or seldom pla-

centas axile. Stylodia free or united.

Stigma is not peltate, sometimes api-

cal or rarely stigmas numerous

(Dendrostigma, Mayna). Ovules

numerous, pendulous, orthotropous

or anatropous, or hemianatropous;

outer integument contributing to the

micropyle. Fruits various, dry or

ﬂ eshy, capsule, drupes, or usually

berries; in Flacourtia the drupes with

separable pyrenes. Seeds exotegmic,

often with thin or thick sarcotesta

(Van Heel 1977), sometimes covered

with long woolly hairs (Calantica) or

with long silky hairs (Bivinia and

Gossypiospermum), winged in

Poliothyrsis, Itoa, and Carrierea;

embryo straight, surrounded by oily,

copious endosperm. Producing cyano-

genic glycosides (Oncobeae, Pangieae,

and occasionally Erythro spermeae and

several species of Banara in Prockieae),

cyclopentenyl fatty acids (‘chaul-

moogra oils’) (Oncobeae, Pangieae,

Azara, Dovyalis, Flacouria in

Flacourtieae) (Spencea and Seigler

1985; Lemke 1988), sometimes pro-

ducing alkaloids, and ﬂ avonols, n =

10–12. . . . . . . . . . 1. flacourtiaceae.

4 Flowers sympetalous, monoecious, solitary

or in fascicles or racemes. Acaulescent

(Guthriea) or climbing (Ceratosicyos) gla-

brous herbs or (Acharia) small subherba-

ceous pubescent shrublets. Vessels mostly

with simple perforations. Fibers with sim-

ple pits, sometimes septate. Leaves alter-

nate, sometimes radical, palmately lobed,

or (Guthriea) unlobed, crenate or serrate,

without stipules. Inﬂ orescences axillary

racemes, fascicles or the ﬂ owers solitary.

Flowers tardily deciduous to deciduous,

small, actinomorphic. Sepals 3–5, free or

adnate to the corolla. Petals united into

3–5-lobed tube, valvate. Stamens 3–5,

attached to the corolla tube near its base or

at the throat; anthers basiﬁ xed, with a

broadly expanded connective, introrse,

opening longitudinally. Staminodes 3–5,

short, ﬂ eshy, internal to the fertile stamens.

Pollen grains 3-colporate. Gynoecium of

3–5 carpels; stylodia united into a more or

less deeply lobed style, stigmas 3–5. Ovary

superior, 1-locular, sessile, with few to

many ovules on parietal placentas. Fruits

3–5- valvate capsules. Seeds with small,

straight embryo and copious, oily

endosperm; raphe expanded (ridged) in

Acharia and Guthriea. Producing cyclo-

pentenoid cyanogenic glucoside and cyclo-

pentenyl fatty acids, gynocardin, ellagic

acid. . . . . . . . . . . . . . . . 11. achariaceae.

3 Petals mostly absent.

11 Stamens numerous. Deciduous trees, gla-

brous or with an indumentum of long sim-

ple hairs. Bark ﬁ ssured, rough, dark brown.

Superorder VIOLANAE 225

Vessels with exclusively scalariform perfo-

rations; lateral pitting opposite. Fibers non-

septate. Xylem with ﬁ bre tracheids

(sometimes septate), without libriform

ﬁ bres. Axial parenchyma apotracheal.

Leaves alternate, large, simple, leathery, dis-

tichous, entire, with two large pits at the base

of the blade on the dorsal surface; stipules

intrapetiolar, free of one another, caducous.

Stomata anomocytic. Inﬂ orescences clusters

of short racemes (Peridiscus), axillary, or at

nodes of fallen leaves, covered with short

branched hairs. Flowers small, actinomor-

phic, fragrant, pale yellow, bisexual, with

large, persistent bracteoles. Sepals 4–5(6) in

Peridiscus or seven in Whittonia, hairy,

imbricate. Petals ﬁ ve (Soyauxia), otherwise

absent Stamens 30–100, seated on or around

the outside of a large, ﬂ eshy, multilobate or

annular disc; ﬁ laments distinct or irregularly

connate toward base, ﬂ exuous (versus

incurved) in Whittonia; anthers 1-locular,

introrse, opening longi tudinally. Pollen

grains 3-colporate, reticulate. Gynoecium of

3–4 carpels; styles 3–4, free, short; ovary

1-locular, superior (half immersed in disc in

Peridiscus), glabrous or lanate (Whittonia);

ovules 6–8, pendulous from the top of a uni-

locular ovary. Fruit drupaceous (known only

from Peridiscus), or (Soyauxia) capsular,

ﬂ eshy, large, 1-seeded. Seeds with a small,

straight embryo lying alongside the copious,

horny endosperm; the cotyledons two, ovate-

lanceolate, membranous. Prismatic crystals

absent. . . . . . . . . . . . . . . . 4. peridiscaceae.

11 Stamen one. Shrubs and small trees. Vessels

with scalariform perforation plates that

have numerous bars. Fibers with simple or

obscurely bordered pits, usually septate.

Rays heterogeneous. Axial parenchyma

diffuse. Leaves alternate, simple, entire or

toothed, distichous, stipules small, cadu-

cous. Stomata more or less distinctly ano-

mocytic. Inﬂ orescences often clustered,

axillary catkinlike dense spikes or looser

racemes. Flowers very small, bisexual or

sometimes polygamous, zygomorphic.

Sepals absent or (1)2–6, unequal; petals

absent. Stamen seated on or within a ﬂ eshy

disc; anther basiﬁ xed, 2-locular, locules

well separated on an expanded connective,

or sometimes individually stipitate, open-

ing longitudinally. Pollen grains 3-colpo-

rate. Gynoecium of 2–3 carpels. Stylodia

united into a style with 2–3 distinct stig-

mas. Ovary 1-locular, superior, with 1–2

ovules on each parietal placenta. Ovules

pendulous, anatropous, with rather thick,

elongate funicle. Fruits 3-valvate capsules,

with three valvular, 1(-3)-seeded. Seeds

with brightly colored, oil-rich sarcotesta,

straight embryo and rather copious, oily

endosperm; cotyledons two, broad and

foliar. . . . . . . . . . . . 3. lacistemataceae.

2 Perianth reduced. Trees or shrubs, sometimes

shrublets (some arctic and subarctic species of

Salix). Roots usually forming ectotrophic mycor-

rhizae. Vessels with simple perforations; lateral

pitting alternate. Fibers with simple pits, occa-

sionally pits with vestigial borders. Rays uniseri-

ate-heterogeneous (Salix and certain species of

the subgenus Turanga of Populus) or homoge-

neous (most species of Populus and Chosenia).

Axial parenchyma scanty, terminal, vasicentric,

or diffuse. Sieve-element plastids of S-type.

Nodes trilacunar. Leaves alternate, simple, entire,

dentate, or lobed, with small or sometimes folia-

ceous stipules. Stomata paracytic. Distal end of

the petiole with one or more closed rings of

xylem and phloem. Flowers small, axillary to a

bract, but without bracteoles, in dense erect

(nearly all species of Salix) or pendulous (Populus

and Chosenia), unisexual, often precocious cat-

kins (spikes or less often racemes with shortened

peduncles), dioecious, anemophilous or (most

species of Salix) secondarily entomophilous.

Calyx vestigial, in Populus more or less disclike

in male ﬂ owers and saucer-shaped or cupulate in

female ones, but in Salix represented by one or

three (rarely 3–5) small nectary glands that may

be united into a lobed ring. In Chosenia, how-

ever, there are usually no glands and female

ﬂ owers are only sometimes provided by two

small lateral glands. Stamens (4)6–40 (rarely up

to 60 or even 70) in Populus 1–2, rarely three or

five (up to 12) in Salix, and 3–6 in Chosenia;

filaments free (Populus

and some spp. of Salix)

or basally connate, rarely connate throughout.

226 Subclass V. DILLENIIDAE

Anthers opening longitudinally. Pollen grains

2-celled or sometimes (in some species of

Populus) 3-celled, usually 3- colporate or

(Populus) inaperturate; in Populus and Chosenia

pollen dry and dispersed by wind, in Salix pollen

viscid and dispersed by insects. Gynoecium of

two, less often of 3–4 (certain species of Populus)

united carpels; stigmas sessile or on a short style,

often biﬁ d or irregularly lobed, carinal (as in

Populus and many species of Salix) or commis-

sural (as in most species of Salix); ovary supe-

rior, sessile or on a short or rarely long gynophore,

unilocular, with numerous (Populus) or 2–10

ovules, in Salix rarely only with one ovule.

Ovules anatropous, weakly bitegmic (some spe-

cies of Populus) or more often unitegmic, crassi-

nucellate. Fruits 2–4-valvate capsules. Seeds

minute, enveloped at the base by a tuft of long

hairs (coma) originating from the placenta; seed

coat thin, formed by the outer integument;

embryo large, straight, with well- developed

cotyledons and hypocotyl and very weakly

developed radicle; endosperm much reduced or

absent. Producing special phenolic heterosides

salicin and populin, only seldom cyanogenic.

n = 11, 12, 19. . . . . . . . . . . . . . . 2 salicaceae.

1. FLACOURTIACEAE

Richard ex A.P. de Candolle 1824 (including

Bembiciaceae R. Keating et Takhtajan 1996, Erythro-

spermaceae Doweld 2001, Gerrardinaceae Alford

2006, Homaliaceae R. Brown 1818, Poliothyrsidaceae

Doweld 2001, Prockiaceae Bertuch 1801, Samydaceae

Ventenat 1808). 82/875. Widespread in the tropics and

subtropics, with some species in temperate regions.

Erythrospermeae: Erythrospermum, Camptostylus,

Ahernia, Dasylepis, Scottellia, Rawsonia; oncobeae:

Mayna, Carpotroche, Grandidiera, Xylotheca, Oncoba,

Caloncoba, Lindackeria, Poggea, Prockiopsis,

Buchnerodendron, Peterodendron; p angieae: Chloro-

carpa, Hydnocarpus (Taraktogenos), Eleutherandra,

Gynocardia, Pangium, Baileyoxylon, Chiangiodendron,

Trichadenia, Scaphocalyx, Ryparosa, Kiggelaria;

homalieae: Dissomeria, Calantica, Bartholomaea,

Homalium, Byrsanthus, Gerrardina, Neopringlea,

Trimeria; bembicieae: Bembicia; scolopieae: Scolopia,

Hemiscolopia, Pseudoscolopia, Phyllo botryon,

Phylloclinium, Mocquerysia; prockieae: Banara,

Pineda, Prockia, Hasseltia, Hasseltiopsis, Macrohasseltia

Neosprucea, Pleuranthodendron; flacourtieae:

Bennettiodendron, Flacourtia, Dovyalis, Tisonia, Azara,

Ludia, Xylosma, Poliothyrsis, Carrierea, Itoa, Idesia,

Olmedlella, Priamosia; aba tieae: Abatia, Aphaerema;

samydeae (casearieae): Casearia (including

Synandrina), Bivinia, Euceraea, Hecatostemon, Osmelia,

Pseudosmelia, Laetia, Neoptychocarpus, Ophiobotrys,

Lunania, Ryania, Samyda, Tetrathylacium, Zuelania,

Lasio chlamys, Trichostephanus.

Closely related to the Berberidopsidaceae, especially

through the most archaic tribe Erythrospermeae, which

is characterized by very primitive wood anatomy as well

as by sepals and petals usually spirally arranged and

often indistinguishable from one another. There are also

similarities with Stachyuraceae (Theales) and

Elaeocarpaceae (Elaeocarpales). A rather heterogeneous

family whose taxonomic boundaries are not quite clear

and have been variously circumscribed. It is especially

difﬁ cult to draw a clear-cut boundary between the

Flacourtiaceae and Passiﬂ oraceae (Passiﬂ orales). The

tribes Paropsieae and Abatieae occupy a somewhat

intermediate position between these two families. They

differ from the typical Flacourtiaceae by the presence of

the extrastaminal corona (weakly developed in the

Abatieae), wood- anatomical (Metcaife and Chalk 1950)

and palynological (Keating 1973) features and stand

nearer to the Passiﬂ oraceae. Lemke (1988) excludes

both of these tribes from the Flacourtiaceae. The correct

position of the Prockieae is somewhat in doubt. Evidence

from xylem anatomy (Miller 1975) and pollen morphol-

ogy (Keating 1973) suggests that the Prockieae are

allied to both the Flacourtiaceae and the Elaeocarpaceae,

thus representing a link between these two families (see

Lemke 1988). Hutchinson (1967) placed the tribe

Prockieae in the Tiliaceae s. 1. (including the

Elaeocarpaceae). According to Miller (1975), a com-

parison of the xylem anatomy of Tiliaceae and

Elaeocarpaceae with the secondary xylem of the tribe

Prockieae suggests a close alliance between the tribe

Prockieae and the Elaeocarpaceae. The family occupies

a basal position in the Violales.

2. SALICACEAE

Mirbel 1815. 3/400. Mainly temperate and cold regions

of the Northern Hemisphere, but also in southern parts

Superorder VIOLANAE 227

of South America, in South Africa, and in Southeast

Asia. Absent in New Guinea and Australia.

Populus, Salix, Chosenia

Very closely related to the Flacourtiaceae, espe-

cially to Idesia and Itoa (Hallier 1908, 1911, 1912;

Holden 1912; Gobi 1916; Cronquist 1957, 1981, 1988;

Takhtajan 1959, 1966, 1987; Thorne 2001), but differs

in unisexual ﬂ owers, reduced perianth, inﬂ orescences

and exotestal seeds. Afﬁ nity with the Flacourtiaceae is

conﬁ rmed by wood anatomy (Gzyrian 1952, 1955;

Miller 1975), pollen morphology (Keating 1973) and

uredinological data (Holm 1969) as well as by chemi-

cal data (salicin in Salicaceae and Idesia and some

other ﬂ acourtiaceous genera).

3. LACISTEMATACEAE

C. Martius 1826. 2/14. West Indies and tropical

America from southern Mexico to southeastern Brazil,

Paraguay, Uruguay, and Argentina.

Lacistema, Lozania

Very close to the Flacourtiaceae, especially to the

Erythrospermeae. Although wood structure is rather

primitive, the ﬂ oral morphology is highly specialized.

4. PERIDISCACEAE

Kuhlmann 1950. 3/11. Northern Brazil and Venezuela

(Peridiscus, 1), Guyana (Whittonia, 1); Soyauxia (9) in

West Africa.

Peridiscus, Whittonia, Soyauxia

Related to the Flacourtiaceae, especially to the

Erythrospermeae.

5. VIOLACEAE

Batsch 1802 (including Alsodeiaceae J.Agardh 1858,

Leoniaceae A.L.P.P. de Candolle 1868). 20/900.

Cosmopolitan, mostly in tropical and subtropical

regions, but with many representatives of the largest

genus Viola (500) in temperate regions. The archaic

genus Rinorea (about 300) is pantropical and espe-

cially characteristic of rain forests.

5.1 VIOLIOIDEAE

Flowers actinomorphic to zygomorphic. Petals

imbricate. Filaments free or more or less connate.

Anthers introrse. Fruits capsules or berries. – rinoreae:

Rinorea, Allexis, Gloeospermum, Melicytus (includ-

ing Hymenanthera), Isodendrion, Rinoreocarpus,

Decorsella, Amphirrhox, Hekkingia, Paypayrola;

violeae: Hybanthus, Agatea, Anchietea, Corynostylis,

Mayanaea (including Orthion), Schweiggeria, Noisettia,

Viola.

5.2 LEONIOIDEAE

Flowers actinomorphic. Petals quincuncial or imbri-

cate. Filaments united into a tube. Fruits nutlike. –

Leonia.

5.3 FUSISPERMOIDEAE

Petals convolute. – Fusispermum.

6. DIPENTODONTACEAE

Merrill 1941. 1/1. Eastern Himalayas, northeastern

India, northern Burma, southeastern Tibet, southwest-

ern China.

Dipentodon.

Despite a free-basal placenta this family is probably

related to the Flacourtiaceae as suggested by

T.A. Sprague (see Metcaife and Chalk 1950: 126).

Pollen grains are ﬂ acourtiacean (Lobreau 1969;

Lobreau-Callen 1982). The perianth of Dipentodon

also resembles that of some genera of the tribe

Homalieae. Dipentodontaceae are rather isolated

within the Violales and Wu et al. (2002) recognized a

separate order Dipentodontales in Dilleniidae.

7. SCYPHOSTEGIACEAE

Hutchinson 1926. 1/1. Borneo.

Scyphostegia.

Probably derived from the Flacourtiaceae (Metcalfe

1956; Takhtajan 1966; Van Heel 1967).

8. PASSIFLORACEAE

A.L. de Jussieu ex Roussel 1806 (including Modeccaceae

Horaninow 1847, Paropsiaceae Dumortier 1829,

Smeathmanniaceae Martius ex Perleb 1838). 17/700–

750. Widely distributed in tropical, subtropical, and

warm-temperate regions, especially in tropical America

and Africa.

228 Subclass V. DILLENIIDAE

paropsieae: Smeathmannia, Barteria, Paropsia,

Paropsiopsis, Androsiphonia, Viridivia;. passifloreae:

Ancistrothyrsus, Passiﬂ ora (including Tetrapathaea),

Hollrungia, Mitostemma, Dilkea, Basananthe (including

Tryphostemma), Schlechterina, Crossostemma, Adenia,

Deidamia, Efulensia.

Derived from the Flacourtiaceae through the archaic

tribe Paropsieae and so close to them that it is difﬁ cult

to draw a clear taxonomic boundary between these two

families. The family Passiﬂ oraceae or its immediate

ancestor was the basal stock from which Turneraceae,

Malesherbiaceae, Caricaceae, and Achariaceae as

well as the order Cucurbitales arose. In contrast to

Flacourtiaceae, which usually produce the dihydroxylated

derivative gynocardin, cyclopentenoid cyanohydrin

glucosides are typical of Passiﬂ oraceae (Jaroszewski

and Olafsdottir 1987).

9. TURNERACEAE

Kunth ex A.P. de Candolle 1828 (including Piriquetaceae

Martynov 1820). 10/150. Subtropical and tropical parts

of America and Africa, Madagascar, and Rodriguez

Island (Mathurina).

Mathurina, Erblichia, Stapﬁ ella, Loewia, Tricliceras,

Hyalocalyx, Streptopetalum, Adenoa, Piriqueta, Turnera.

Very close to the Passiﬂ oraceae and Malesherbiaceae

and produce structurally related cyanogens. The

Turneraceae seem to be similar to the Malesherbiaceae

and the primitive members of the Passiﬂ oraceae in pro-

ducing only four basic cyclopentenoid rings (Spencer

et al. 1985). But their ﬂ owers have no extrastaminal

corona (very narrow corona only in Piriqueta and

Erblichia), and their petals are contorted in the bud.

10. MALESHERBIACEAE

D. Don 1827. 1/24. Western South America: Paciﬁ c

coastal desert and adjacent arid Andes of Chile (includ-

ing Atacama), Peru, and Argentina.

Malesherbia.

Very close to the Passiﬂ oraceae and Turneraceae

and have structurally related cyanogens and possess

similar hydrolytic enzymes (Spencer and Seigler

1985). Differ mainly in contorted petals. They have

both extrastaminal corona and androgynophore, but

seeds are without arils.

11. ACHARIACEAE

H. Harms 1897. 3/3. Warm-temperate South Africa

(Cape Province, centered in the eastern Cape).

Acharia, Ceratosicyos, Guthriea.

Closely related to the Passiﬂ oraceae, which is sup-

ported also by chemical data (the presence of the

cyclopentenoid cyanogenic glucoside and gynocar-

din). From the Passiﬂ oraceae they differ mainly in

sympetalous corolla (“a feature not often associated

with a unisexual condition, except in Cucurbitaceae

and in a few genera of Euphorbiaceae,” according to

Hutchinson [1969: 221]).

12. APHLOIACEAE

Takhtajan 1985. 1/1–4–8. Tropical East Africa, Madagascar,

Comoros, Seychelles, Mascarene Islands.

Aphloia (Neumannia).

Rather isolated family, which probably related to

Flacourtiaceae.

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Order 67. ELAEOCARPALES

Trees or shrubs, pubescent with unicellular or glandu-

lar hairs. Mucilage cavities and canals absent, but

mucilage cells are known to occur. Vessels usually

with simple and oblique perforation plates, but some

vestigial scalariform perforations with a few bars occur

in Aristotelia, Vallea and Tremandra): lateral pitting

large, typically opposite, occasionally alternate or

transitional to alternate in some genera. Fibers with

small bordered pits (the borders sometimes very incon-

spicuous) or with simple (Platytheca and Tremandra)

pits, mostly septate. Rays markedly heterogeneous,

mixed uniseriate and pluriseriate, the latter often with

long ends. Axial parenchyma paratracheal or some-

times absent, in Tremandreae diffuse to vasicentric, in

narrow sheaths around the vessels, conﬂ uent, and dif-

fuse in Echinocarpus. Phloem not stratiﬁ ed. Sieve-

element plastids of S-type. Nodes trilacunar or

unilacunar with one trace. Cells of the epidermis

frequently mucilaginous. Leaves alternate or sometimes

opposite, simple, entire, crenate, serrate, or denticu-

late, stipulate or very rarely estipulate; stipules persis-

tent or caducous. Stomata paracytic, encyclocytic, or

(Tremandreae) anomocytic. Flowers in racemes, pani-

cles, or in dichasia, rarely solitary in the axils, bisexual

or sometimes polygamous, actinomorphic, often apet-

alous. Sepals 4–5, free or more or less connate, valvate

or less often imbricate. Petals 4–5, free or rarely basally

connate, rarely united high up into a corolla crenate-

dentate at the apex, entire, lobed, dentate, incised, or

laciniate, valvate or seldom imbricate, sometimes

wanting. Annular extrastaminal nectary disc with lobes