Takhtajan Armen. Flowering Plants

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132 Subclass IV. CARYOPHYLLIDAE

somewhat ﬂ attened, erect-spreading, very

shortly connate at the base, the inner side papil-

late; ovule solitary. Fruits woody, 2-locular and

1–2-seeded capsules. Seeds partially enclosed

by a yellow aril.. . . . . . . . . 5. barbeuiaceae.

8 Petioles not articulated. Plants not blackening

on drying.

9 Seeds covered by a large red aril. Small, gla-

brous trees or shrubs, usually with decum-

bent or scandent branches. Leaves alternate,

entire, ﬂ eshy, with trans lucent margins; dif-

fuse axial parenchyma presents (Carlquist

1999). Crystalloids within the sieve-element

plastids polygonal. More or less scandent or

sprawling glabrous shrubs. Flowers in termi-

nal racemes. Pedicels bracteate and bracteo-

late at the base. Sepals ﬁ ve, imbricate,

persistent in the fruit. Petals free, imbri-

cate, shorter than the sepals. Stamens ten,

in one series; ﬁ laments connate at the base;

anthers dorsiﬁ xed, cordate, introrse. Pollen

grains 3-colpate, operculate, with spinu-

lose and punctate rectum. Ovary superior,

1- locular, with a very short style and 3–5

subulate recurved stigmas, papillose inside.

Ovules as many as the stigmas, amphitrop-

ous, basal around a central column. Fruits

coriaceous, 3–5-valved and 1–5-seeded

capsule. Seeds almost covered by a large

red aril, exotesta more or less palisade,

unligniﬁ ed, endotegmen enlarged, persis-

tent, n = 36 (P. Goldblatt, 2003). . . . . . . . .

9. stegnospermataceae.

9 Seeds without aril, or when an aril is pres-

ent, it does not cover the seed.

10 Fruits 1-seeded berries. Shrubs or small

trees with normal secondary growth.

Leaves alternate, entire, estipulate.

Flowers in axillary racemes or panicles,

dioecious. Sepals 4–5. Stamens 10–20;

ﬁ laments slender, free or basally con-

nate. Pollen grains with mostly 4–6 large

pores or more or less irregular apertures,

which are often poorly deﬁ ned.

Gynoecium of two carpels with short,

basally connate stylodia; ovary superior,

1-locular, with 1 (very rarely 2) campy-

lotropous basal ovule. Seeds black,

tuberculate, with very small aril at the

hilum. Present C-glycosylﬂ avonoids.. . .

6. achatocarpaceae.

10 Fruits dry.

11 Fruits dehiscent.

12 Stamens opposite the sepals. Annual

herbs to small trees with normal

secondary growth. Leaves alternate

or sometimes opposite, often with

hairs, bristles, or scales in the axils;

stipules scarious. Flowers bisexual

or rarely unisexual, with two or

sometimes ﬁ ve or more (Lewisia)

imbricate bracteoles and (2)5(-18)

free or basally connate, more or less

petaloid and often caducous sepals.

Stamens commonly as many as and

opposite the sepals. Pollen grains

ranging from 3-colpate to pantocol-

pate and pantoporate. Gynoecium

2–8-carpellate, with more or less

united stylodia and capitate stig-

mas. Ovary superior to inferior,

with two to many or rarely one

ovule; ovules anatropous, acampy-

lotropous, or amphitropous, placen-

tation free-central or basal. Fruits

circumscissile or valvate capsules,

rarely nutlike. Seeds sometimes

arillate; embryo curved to annular, n

= 8–12 (-17), mostly 8 or 9, in

Claytonia virginiana n = 6–95 (Bogle

1969).. . . . . . 10. portulacaceae.

12 Stamens alternate with the petals.

Densely caespitose, glabrous sub-

shrubs; cuticle waxes as procumbent

platelets. Leaves alternate, imbri-

cated all around the stem, without

stipules. Flowers solitary, axillary,

with (1)2(3) lateral, apical or basal

bracteoles and 4–5(6) petaloid sepals.

Stamens 3–5(6), free or adnate to

calyx tube when it is present. Pollen

grains 3-colpate. Gynoecium

2-carpellate, unilocular; ovary with

4–7 ovules borne near the base of

central column. Fruits 1–5-seeded

capsules, or indehiscent. Seeds with

endosperm and curved embryo,

n = 48. . . . . . 11. hectorellaceae.

Superorder CARYOPHYLLANAE 133

11 Fruits indehiscent.

13 Sepals united to form a lobed and

commonly corolloid tube and some-

times subtended by brightly colored

sepaloid bracteoles. Trees, shrubs,

herbs, and lianas, sometimes thorny.

Nodes swollen. Wood storied, ray-

less. Vessels with simple perfora-

tion, occasionally also reticulate;

stem with anomalous secondary

growth. Leaves opposite or rarely

alternate, toothed. Inﬂ orescences

various, cymose, very rarely a

raceme; ﬂ owers bisexual or rarely

dioecious (Cephalotomandra).

Tubular, petaloid perianth with ind-

uplicate-valvate or contorted aesti-

vation. Stamens one to many, free or

connate at the base, involute in bud;

ﬁ laments often of unequal length;

anthers dorsiﬁ xed near the base.

Pollen grains 3(4)-colpate, pericol-

pate, or periporate. Ovary superior,

1-locular, with slender styles. Ovule

solitary, bitegmic, or rarely uniteg-

mic (Abronia umbellate, Boerhaavia

diffusa). Fruits nutlike, surrounded

by accrescent (viscid) basal part of

the perianth; endotesta thickened

(Mirabilis), testa multiplicative;

embryo peripheral, arcuate or rarely

straight, green; perisperm copious

to scanty; n = 8–11(13), rarely 17,

mostly 10. . . . .7. nyctagina ceae.

13 Sepals not forming a corolloid

calyx tube.

14 Perennial herbs with slender,

annual twining or scrambling

stems from ﬂ eshy-thickened or

tuber- bearing rhizomes; vascu-

lar bundles separate, bicollateral.

Leaves opposite, conduplicate

(Anredera). Flowers in race-

mose inﬂ orescences, small,

mostly bisexual. Bracteoles

two, often united to the base of

the calyx. Sepals ﬁ ve, often col-

ored, almost free or connate

below, persistent. Stamens ﬁ ve,

opposite to the sepals and adnate

to the base of sepals or to the

calyx tube. Pollen grains cuboid,

pantocolpate. Gynoecium of

three carpels; ovary unilocular

when mature, with three stylo-

dia or a style, often 3-lobed.

Ovule solitary, basal, anatro-

pous to campylotropous. Fruits

indehiscent, surrounded by

persistent, often ﬂ eshy calyx or

by winglike persistent bracte-

oles; exotesta and endotesta

more or less thickened, embryo

green, starch grains clustered;

n = 12, 22.. . 12. basellaceae.

14 Not twining or scrambling

annual herbs. Flowers unisexual,

monoecious. Bracteoles two,

very unequal. Sepals in male

ﬂ owers four, free, membranous,

in female ﬂ owers lacking.

Stamens four, alternating with

sepals; ﬁ laments slender, long.

Pollen grains cuboid, 6-porate.

Gynoecium of three carpels;

ovary unilocular, with three free

stylodia. Ovule solitary, basal,

evidently campylotropous.

Fruits nutlets, several becoming

enclosed by inﬂ orescence axis,

n = 12. . . 13. halophytaceae.

6 Ovary mostly inferior or semi-inferior.

15 Ovary with a single fertile locule and 2(3) empty

locules. Very spiny, cactuslike, xerophytic

shrubs and trees with soft wood and rather well-

developed pith; cuticular waxes as ribbons or

rodlets. Leaves alternate, entire, small, some-

times much reduced. Flowers mostly unisexual

and usually dioecious. Sepals four, in two series.

Petals absent. Stamens 8(10), in two cycles

basally connate and with adaxial nectaries.

Pollen grains 3-celled, (4)5–7-zonocolpate.

Gynoecium of (2)3(4) carpels with a style and a

usually irregularly (2)3(4)-lobed, peltate,

fringed stigma; ovary with as many locules as

carpels, but usually only one locule is fertile

and bears a solitary, basal ovule. Fruits dry,

indehiscent, usually 3-angled. Seeds with a

134 Subclass IV. CARYOPHYLLIDAE

small funicular aril, nearly or quite without both

perisperm and endosperm. Present methylated

ﬂ avonoids; n = 8.. . . . . . . . . 15. didiereaceae.

15 Ovary without empty locules.

16 Plants leaf-succulents or rarely stem-succu-

lents, always unarmed. Leaves alternate or

opposite, usually entire, rarely toothed.

Stomata anisocytic, anomocytic or para-

cytic. Nodes 3-locular with three traces.

Anomalous secondary growth present.

Epicuticular waxes present. Inﬂ orescences

axillary or terminal pleiochasia, dichasia or

ﬂ owers solitary. Flowers cyclic, bisexual or

rarely unisexual (Glischrothamnus). Calyx

lobes 5–8; petals ca. 30–40, or absent.

Stamens (1-)4–5, or 8–10, or numerous.

Pollen grains 3-colpate or rarely 3-colpo-

rate. Ovary superior, half-inferior, or infe-

rior, surrounded by nectary ring. Female

gametophyte usually Polygonum-type.

Ovules campylotropous or anacampylotro-

pous, funicle often long. Fruits usually locu-

licidal capsules, rarely baccate (Carpobrotus),

sometimes nutlike. Seeds often arillate.

n = 8, 9, mostly 9. . . . . . . . . 8. aizoaceae.

16 Plants stem-succulents or with distinctly

spiny stem, cork also in cortex; rays wide and

tall; nodes often with two or many traces;

mucilage copious; cuticular waxes as ribbons

or rodlets. Leaves mostly very reduced.

Stomata usually paracytic. Flowers usually

solitary, rarely in panicles, spiral or spirocy-

clic, bisexual, rarely unisexual. Perianth seg-

ments spirally arranged, usually not

differentiated into sepals and petals, Stamens

numerous, centrifugal; anthers small, dorsi-

ﬁ xed or basiﬁ xed. Pollen grains 3-colpate to

6–15-colpate or -porate. Gynoecium of

3-many carpels; stigma wet, lobed; ovary

inferior or rarely (in spp. of Pereskia) supe-

rior, placentation more or less parietal; ovules

numerous, anatropous to campylotropous.

Fruits usually indehiscent, commonly ﬂ eshy

and baccate, rarely dry, funicles ﬂ eshy in

fruit; endotegmic cell walls thickened or not;

n = 11.. . . . . . . . . . . . . . . . . .14. cactaceae.

3 Producing anthocyanins but not betalains.

17 Leaves opposite to alternate or verticillate,

slightly or not succulent; stipules scarious, small

or absent.

18 Seeds small, dark brown to shiny black.

Herbs, subshrubs, rarely shrubs with anoma-

lous secondary growth. Inﬂ orescences pleio-

chasail or dichasial cymes, or in glomerules

to cincinnii, or ﬂ owers solitary. Sepals ﬁ ve,

mostly free, rarely shortly connate. Petals

small and inconspicuous or more often want-

ing. Stamens 5–10 to many, ﬁ laments ﬁ li-

form, free or shortly connate; anthers

dorsiﬁ xed; nectary disc absent or annular.

Ovary (except in Adenogramma) with two to

several locules, at least below, but in the upper

part of the ovary the partitions sometimes

incomplete. Placentation axile (except

Adenogramma). Fruits usually loculicidal or

circumscissible capsules, often surrounded

by persistent calyx. Seeds sometimes arillate,

embryo curved or around the periphery.

Plants containing anthocyanins; n = 8

(Hypertelis) or 9. . . . . 16. mollugina ceae.

18 Seeds rather large, pale yellow. Herbs or

subshrubs, glabrous or with glandular

hairs. Sieve-element plastids of PIIIc’’f-

type. Leaves alternate to subopposite, lin-

ear, subulate, lanceolate or orbicular,

estipulate. Inﬂ orescence cymose, lax or

glomerulate. Flowers small, sepals ﬁ ve,

petals ﬁ ve or absent, adnate to the stami-

nal tube. Stamens (5)-7(-10). Pollen

grains tri colpate. Ovary pseudomonomer-

ous 2-chambered, with one ovule per

chamber. Fruit separating into two woody

cocci; rarely broadly winged; seeds with

rough surface, strictly reniform, slightly

winged, periclinals ﬂ at, micropapillate,

anticlinals highly raised, laciniate or with

longitudinal striations (Hassan et al.

2005), testa with cells in rows along the

dorsal junction; n = 9.. . . 17. limeaceae.

17 Leaves opposite or rarely (as in Telephium

and Corrigiola) alternate; stipules absent or

present (Illecebroideae), and then often

scarious, white or brownish. Stomata often

diacytic or anisocytic. Mostly herbs, peren-

nial or annual, or sometimes subshrubs, very

rarely (Sanctambrosia manicata) small

trees; stems often swollen at the nodes due

to an anomalous growth of the concentric

rings of xylem and phloem. Inﬂ orescences

usually terminal, paniculate, racemiform or

Superorder CARYOPHYLLANAE 135

capitate dichasial cymes, or ﬂ owers solitary.

Flowers actinomorphic, or very rarely

(Mediterranean Drypis spinosa) slightly

zygomorphic, bisexual or rarely the plants

dioecious. Sepals (4)5–25, rarely spirally

arranged, free or connate. Petals mostly more

or less developed, in Illecebroideae often

absent. Stamens 10 (3–13), ﬁ laments ﬁ liform,

free, anthers dorsiﬁ xed. Gynoecium of 2–5(-

10) carpels; ovary superior, unilocular above,

but usually more or less distinctly partitioned

toward the base, at least when young; ovules

mostly numerous, but sometimes few or even

solitary. Fruits capsules, dehiscent by apical

teeth or by valves, rarely circumscissile, or

rarely a scarcely ﬂ eshy berry (Cucubalus).

Seeds not arillate (except seeds with funicu-

lar elaiosome in Moehringia), embryo periph-

eral, arcuate to spiral (Dripis), or straight

(Dianthus, Stellaria). Produce anthocyanins

(batalains) and anthraquinones, saponins,

ferula acid and pinitol, n = 5–15, 17–19,

mostly 8. . . . . . . . . . 18. caryophylla ceae.

2 Sieve-element plastids without protein crystal.

19 Stylodia three. Xerophytic, evergreen, freely

branched, shortly pubescent shrubs or small trees

with anomalous secondary growth of concentric

type (concentric rings of vascular bundles); hairs

uniseriate. Vessels with simple perforations and

with tertiary thickenings; lateral pitting alternate.

Fibers with large bordered pits. Axial parenchyma

apotracheal, very rare, except for conjunctive tis-

sue associated with the anomalous structure.

Sieve-element plastids of Ss-type. Nodes unilacu-

nar. Leaves small, opposite, sessile, simple but

jointed at the base, entire, pinnately veined, leath-

ery, estipulate; the mesophyll containing calcium

oxalate crystals. Stomata laterocytic or cyclocytic,

or anomocytic (and also of intermediate type).

Flowers unisexual, dioecious, actinomorphic,

apetalous. Male ﬂ owers small, in cernuous, capit-

ulate, axillary clusters, the female ones larger, and

mostly solitary in the axils (more rarely in pendu-

lous 2–7-ﬂ owered racemes). Sepals (4)5(6), much

imbricate, ﬁ mbriate, accrescent in female ﬂ ow-

ers. Stamens (8)10(12), free, inserted more or less

distantly on ﬂ at receptacle; ﬁ laments short and

stout; anthers elongate, tetrasporangiate, bas-

iﬁ xed, 2-locular, extrorse, opening longitudinally.

Pollen grains 2-celled, with three large, weakly

deﬁ ned, porelike apertures the membranes of

which are bordered with irregularly shaped insu-

lae or granula forming an operculoid structure in

the central part and grading to the exine pattern.

Pistillodia absent. Gynoecium of three united car-

pels, with three elongate, reﬂ exed, subulate, papil-

late, hairy, deciduous stylodia clustered at the top.

Ovary superior, 3-locular, with one pendulous,

apical-axile ovule per locule. Ovules anatropous,

or apotropous, pendulous, bitegmic, crassinucel-

late (but tenuinucellate, according Köhler 2003).

Endosperm nuclear and very reduced. Fruits

ovoid loculicidal capsules, with shiny brown

coriaceous pericarp and usually one seeded by

abortation (two of the locules empty), rarely

2–3-seeded. Seeds large, subtriquatrous, red-

brown, glandular, and short-haired. Embryo

straight, with thick cotyledons; cotyledons and

peripheral layers of the hypocotyl contain a

cyanogenic glucoside (simmondsin), mono-

ethylenic acids, and a high proportion of a

unique liquid wax; endosperm scanty or wanting,

n = 13.. . . . . . . . . . . . . . . . .19. simmondsia ceae.

19 Stylodium one.

20 Sepals usually dry and scarious or membra-

nous, sometimes absent. Herbs, rarely shrubs

or small trees. Vessels with simple perfora-

tion. Leaves alternate or opposite, entire,

estipulate. Inﬂ orescences terminal and axil-

lary spikes, heads, racemes or cymes.

Stamens often connate at the base into a

tube. Stylodia united into a style with 2–3-

ﬁ d or capitate stigma. Epicuticular wax struc-

tures lacking, n = 7–11, 13, 15, 17, mostly

8 and 9. . . . . . . . . . . . .20. amaranthaceae.

20 Sepals mostly green or greenish and usually

more or less herbaceous or somewhat mem-

branous, rarely wanting. Herbs and shrubs,

rarely small trees or climbers. Vessels with

simple perforation or very rarely with some

oblique scalariform perforations (Axirys).

Leaves mostly alternate, rarely opposite,

sometimes ﬂ eshy or reduced to scales,

entire or toothed, estipulate. Stamens free

or sometimes connate at the base. Stylodia

distinct or more or less connate into a

style. Female gametophyte Polygonum- or

Allium-type. Epicuticular wax structures

present (more or less lobed platelets), n = 9,

rarely 6. . . . . . . . . . 21. chenopodiaceae.

136 Subclass IV. CARYOPHYLLIDAE

1. PHYTOLACCACEAE

R. Brown 1818 (including Agdestidaceae Nakai 1942,

Hilleriaceae Nakai 1942, Lophiocarpaceae Doweld et

Reveal 2005, Petiveriaceae C. Agardh 1824, Rivina-

ceae C. Agardh 1824, Sarcocaceae Raﬁ nesque 1837,

Seguieriaceae Nakai 1942). 15/50. Subcosmopolitan.

1.1. PHYTOLACCOIDEAE

Fibers vasicentric. Perianth segments usually 5.

Carpels 3–17, free or united. Fruit a berry. – Anisomeria,

Ercilla, Phytolacca, Nowickea.

1.2. RIVINIOIDEAE

Shrubs, lianas or herbs. Perianth segments usually 4.

Carpel one. Fruits indehiscent, often samaroid, pericarp

mostly adherent to the seed. – rivineae (petiver-

ieae): Rivina, Trichostigma, Schindleria, Hilleria,

Ledenbergia, Petiveria, Monococcus; seguierieae:

Gallesia, Seguieria; lophiocarpeae: Lophiocarpus.

1.3. AGDESTIDOIDEAE

Semiwoody liana with massive globose rootstock and

anomolous secondary growth. True tracheids present.

Sepals four, rarely ﬁ ve (in terminal ﬂ owers). Carpels

(3)4, united. Ovary semi-inferior. Fruits a 1-seeded

achene with sepalline wings. – Agdestis.

2. RHABDODENDRACEAE

Prance 1968. 1/4. Tropical South America (the Guianas

and Brazil).

Rhabdodendron.

Prance (1968, 1972) closed Rhabdodendraceae

with Phytolaccaceae. Gadek et al. (1996) suggested a

relationship Rhabdodendraceae with the caryophyllid

clade in which Phytolaccaceae are also placed based

on molecular evidence. Fay et al. (1997) indicated that

Rhabdodendron belongs within Caryophyllidae, but

without a close relationship to other families. APG II

(2003) includes Rhabdodendraceae in to Caryophylales

sensu lato. The family is very isolated and represent a

separate order and even superorder (Doweld 2002).

3. GISEKIACEAE

Nakai 1942. 1/7. From tropical and South Africa to

South Vietnam, but mainly African.

Gisekia.

Beginning with Bentham and Hooker (1867), many

authors assigned Gisekia to the Aizoaceae. Others,

including Cronquist (1981), Thorne (1983, 2005) and

Rohwer (1993), transfer it to the Phytolaccaceae s. 1.

Hutchinson (1926, 1959, 1973) includes Gisekia in the

Molluginaceae, with some members of which, espe-

cially Limeum, it shares some morphological features.

Embryological data also show a close relationship of

Gisekia with Mollugo (Raghavan and Srinivasan 1940).

However, Gisekia differs from the Molluginaceae not

only in its apocarpous gynoecium and indehiscent fruits

but also in the presence of betalains. From the Aizoaceae,

with which Gisekia shares some chemical characters

(Narayana and Narayana 1988; Richardson 1981), it

differs in its apocarpous gynoecium as well as its free

sepals, apetalous ﬂ owers, fruit morphology, etc. From

the Phytolaccaceae it differs in cymose inﬂ orescences

and dry, nutlike fruitlets as well as in the morphology of

its vegetative organs. However, the endexine of Gisekia

suggests a relationship to the Phytolaccaceae (Nowicke

in Behnke and Mabry 1994).

4. SARCOBATACEAE

Behnke 1997. 1/2. North America.

Sarcobatus.

The genus Sarcobatus are usually including in the

Chenopodiaceae. However, according to Behnke (1997),

Sarcobatus markedly differs from Cheno podiaceae. The

important feature that sets off Sarcobatus from the

remainder of the Chenopodiaceae are speciﬁ c sieve-ele-

ments plastids. The sieve-element plastids of Sarcobatus

are of the form P3cf with a globular central and an addi-

tional rodlike protein crystal not found elsewhere in the

Chenopodiaceae (Behnke 1993). “If I had to decide by

the sieve-element plastid data where to place Sarcobatus”

says Behnke, “I would select the suborder Phytolacineae,

where some genera of the Phytolaccaceae and

Nyctaginaceae have the same characteristics.”

5. BARBEUIACEAE

Nakai 1942. 1/1. Madagascar.

Barbeuia.

This large woody vine may be regarded as a very

advanced and specialized derivative of the Phytol-

accaceae. Sieve-element plastids are similar to those of

the Phytolaccaceae (Behnke 1993).

Superorder CARYOPHYLLANAE 137

6. ACHATOCARPACEAE

Heimerl 1934. 2/10. From Texas and northwestern

Mexico to Paraguay and Argentina.

Achatocarpus, Phaulothamnus.

Probably derived from the Phytolaccaceae. Among

the many differences from the Phytolaccaceae are the

sieve-element plastids with polygonal central crystals

(Behnke in Behnke and Mabry 1994).

7. NYCTAGINACEAE

A. L. de Jussieu 1789 (including Allioniaceae

Horaninow 1834, Bougainvilleaceae J. Agardh 1858,

Mirabilidaceae W. R. B. Oliver 1936, Pisoniaceae J.

Agardh 1858). 32/400. Tropical and subtropical

regions, particularly in America. Phaeoptilum is

endemic to southwestern Africa.

boldoeae: Boldoa, Salpianthus, Cryptocarpus;

leucastereae: Andradea, Ramisia, Leucaster,

Reichenbachia; nyctagineae: Colignoma, Pisoniella,

Boerhavia; Anulocaulis, Cyphomeris, Commicarpus,

Caribea, Acleisanthes, Selinocarpus, Okenia, Mirabilis,

Cuscatlania, Nyctaginia, Allionia, Phaeoptilum;

abronieae: Abronia, Tripterocalyx; bougainvilleeae:

Bougainvillea, Belemia; pisonieae: Pisonia, Guapira,

Neea, Neeopsis, Cephalotomandra, Grajalesia.

An advanced entomophilous family derived from

the Phytolaccaceae stock. Stands close to the

Petiveriaceae. According to Baillon (1875), the gynoe-

cium of Nyctaginaceae is constructed like that of the

Rivineae.

8. AIZOACEAE

Martynov 1820 (including Galeniaceae Raﬁ nesque

1819, Mesembryaceae Dumortier 1829, Mesembryan-

themaceae Burnett 1835, Sesuviaceae Horaninow

1834, Tetragoniaceae Link 1831). Circa 130/2500.

Dryer parts of tropical and subtropical regions, but

centered mainly in South and southwestern Africa;

there is also a small center in western and southern

Australia.

8.1 AIZOOIDEAE

Annuals to perennials, with herbaceous or woody

branches; leaves slightly succulent, mostly alternate,

ﬂ at or more rarely cylindrical; inﬂ orescence leafy,

perianth internally petaloid, externally sepaloid; ﬂ ow-

ers perigynous, epigynous, or rarely hypogynous, with

a holonectary; capsules loculicidal or septicidal, often

hygrochastic or a winged nut or horn-shaped, n = 8. –

Gunniopsis, Aizoanthemum, Aizoon, Plinthus, Galenia,

Acrosanthes.

8.2 TETRAGONIOIDEAE

Shrublets or perennials with alternate, entire leaves.

Flowers small, inconspicuous, in andromonoecious

inﬂ orescence. Fruits nutlike, usually with acute horns,

n = 8. – Tetragonia, Tribulocarpus.

8.3. SESUVIOIDEAE

Prostrate to erect perennials or annuals; leaves slightly

succulent, petiolate, often stipulate, ﬂ at or more

rarely cylindrical; inﬂ orescence bracteate; perianth

internally petaloid, externally sepaloid; ﬂ ower perig-

ynous, with a holonectary; fruit a circumscissile cap-

sule or compound and fused to spinous bracts; seed

arilate, n = 8. – Cypselea, Sesuvium, Trianthema,

Zaleya.

8.4 MESEMBRYANTHEMOIDEAE

Annual to short-lived or perennial herbs or shrubs, cor-

tex with vascular bundles; leaves succulent, usually

ﬂ at to almost cylindrical and channeled, bladder cells

often large and conspicuous; ﬂ owers with petals of sta-

minodial origin, with a central placenta, ovary half-

inferior or inferior, nectaries koilomorphic, i.e., always

separated and grooved; fruit a hygrochastic capsule

with purely septal expanding keels reaching from the

central columella to the tip of the valve or very rarely

a woody nut; n = 9. – Mesembryanthemum,

Synaptophyllum, Phyllobolus, Aptenia, Aspazoma,

Aridaria, Caulipsolon, Prenia, Sceletium, Psilocaulon,

Brownanthus.

8.5 RUSCHIOIDEAE

Perennial shrubs, rarely annual; leaves succulent, usu-

ally cylindrical or trigonous, rarely ﬂ at, epidermis

rarely with conspicuous bladder cells, usually xero-

morphic; ﬂ owers with petals of staminodial origin,

with a basal or pariental placenta, ovary inferior, nec-

taries lophomorphic, i. e., crested, either separated or

in a ring or rarely glands ﬂ attened; fruit a hygrochas-

tic capsule with expanding keels of mainly valvar ori-

gin with only small septal portion near the outer rim

of the locule, never reaching to the center of the fruit,

138 Subclass IV. CARYOPHYLLIDAE

mostly with covering membranes and additional clos-

ing devises; n = 9. – apatesieae: Apatesia,

Carpanthea, Hymenogyne, Caryotophora, Conicosia,

Suphesia, Skiatophytum; dorotheantheae:

Aethephyllum, Cleretum, Dorotheanthus; RUSCHIAE:

Bergeranthus, Bijlia, Carruanthus, Cerochlamys,

Chas matophyllum, Corpuscularia, Delosperma, Drosan-

themum, Ecto tropis, Faucaria, Frithia, Gib baeum,

Hereroa, Machairophyllum, Malephora, Mestoklema,

Mossia, Muiria, Neohenricia, Orthopterum,

Oscularia, Rabiea, Rhinephyllum, Rhombophyllum,

Ruschianthus, Trichodiadema, Stomatium, Deilanthe,

Smicrostigma, Oophytum, Vanzijlia, Schlechteranthus,

Nananthus, Hammeria, Marlothistella, Zeuktophyllum,

Acrodon, Didymaotus, Namibia, Wooleya, Aloinopsis,

Dinteranthus, Juttadinteria, Ihlenfeldtia, Cheiridopsis,

Diplosoma, Jacobsenia, Sarcozona, Octopoma,

Vlokia, Pleiospilos, Titanopsis, Osammophora,

Amphibolia, Circandra, Antegibbaeum, Disphyma,

Cylindro phyllum, Ottosonderia, Scopelogena, Hart-

manthus, Dicrocaulon, Leipoldtia, Arenifera,

Meyerophytum, Dracophilus, Feenestraria, Eber-

lanzia, Glottiphyllum, Jordaaniella, Odontophorus,

Ebrac teola, Argyroderma, Enarganthe, Hallianthus,

Stayneria, Khadia, Lapidaria, Nelia, Lithops, Astridia,

Mitrophyllum, Braunsia, Jensenobotrya, Conophytum,

Lampranthus, Carpobrotus, Erepsia, Ruschian-

themum, Ruschia, Monilaria, Namaquanthus,

Antimima, Vanheerdea, Schwantesia, Tanquana,

Cephalophyllum, Stroeberia.

Closely related to the Phytolaccaceae, but more

advanced.

9. STEGNOSPERMATACEAE

Nakai 1942. 1/4. From Baja California to Central

America and West Indies.

Stegnosperma.

Stegnosperma is usually included in the

Phytolaccaceae, from which it differs in its ﬂ ower

morphology (Hofmann 1973), particularly in the pres-

ence of petaloid staminodia as well as in sieve-element

plastids with polygonal protein crystal as in

Achatocarpaceae and Caryophyllaceae (Behnke 1976b,

1982; Behnke in Behnke and Mabry 1994), in seeds

almost completely covered by a large ﬂ eshy funicular

aril, in capsular fruits, and in the presence of quercetin

and kaempferol glycosides (Richardson 1981). They

differ also in wood anatomy. According to Carlquist

(1999), diffuse axial parenchyma is common in

Stegnosperma, but has not been reported in other

Caryophyllales. He also agrees with Rodman (1994),

that Stegnosperma is basal in Caryophyllales.

10. PORTULACACEAE

A.L. de Jussieu 1789 (including Montiaceae

Raﬁ nesque 1820, Talinaceae Doweld 2001). 28/450 or

more. Very widely distributed, but mainly in warm

and temperate regions, particularly well represented

in South Africa, western North America and southern

South America.

portulaceae: Portulaca, Grahamia; talineae:

Talinopsis, Talinaria, Anacampseros, Lewisia, Calan-

drinia, Montiopsis, Erocallis, Calandrinia (including ?

Baitaria), Cistanthe, Rumicastrum, Monocosmia,

Talinum, Talinella, Amphipetalum, Schreiteria, Lenzia,

Calyptridium (including Spraguea), Claytonia, Lim-

nalsine, Naiocrene, Neopaxia, Mona, Montia, Monti-

astrum, Maxia, Crunocallis.

Allied to the Aizoaceae and Cactaceae and espe-

cially to the Hectorellaceae and Basellaceae. Probably

derived directly from the Phytolaccaceae stock.

11. HECTORELLACEAE

Philipson et Skipworth 1961. 2/2. New Zealand (South

Island) and Kerguelen Island.

Hectorella, Lyallia.

Very close to the Portulacaceae, from which they

differ in stamens alternating with the sepals and attached

to the calyx tube and in the densely imbricate arrange-

ment of coriaceous estipulate leaves. Sieve-element

plastids with globular protein crystal resemble those of

the Portulacaceae (Behnke 1975, 1976b, 1993).

12. BASELLACEAE

Raﬁ nesque 1837 (including Anrederaceae J. Agardh

1858 and Ullucaceae Nakai 1942). 4/20. Mainly tropi-

cal and subtropical regions of America, but also tropi-

cal Africa, Madagascar, southern Asia, New Guinea,

and some Paciﬁ c islands.

Superorder CARYOPHYLLANAE 139

baselleae: Basella, Ullucus, Tournonia; boussin-

gaultieae: Anredera (including Boussingaultia).

Very close to the Portulacaceae, especially to Montia,

with which they share not only similar features of ﬂ oral

morphology but also the presence of bicollateral bundles

resulting from the tardy development of internal phloem,

the absence of anomalous secondary growth, the pres-

ence of mucilaginous cells as well as sieve-element

plastids with central globular protein crystal (Behnke

1976b) and the absence of epicuticular wax projections

(Engel and Barthlott 1988; Barthlott in Behnke and

Mabry 1994). However, from the Portulacaceae the

family Basellaceae differs not only in the twining or

scrambling habit but also in its considerably larger pro-

tein crystals of the sieve-element plastids (Behnke in

Behnke and Mabry 1994), solitary ovules (among the

Portulacaceae only Portulacaria and Ceraria have a

solitary ovule), and its indehiscent fruit (indehiscent

nutlike fruits are characteristic of the somewhat transi-

tional genus Portulacaria). The family is divided into

two well- deﬁ ned groups – Baselleae, with ﬁ laments

erect and straight in the bud and spirally twisted embryo,

and Boussingaultieae, with ﬁ laments reﬂ exed outward

in the bud and an annular embryo.

13. HALOPHYTACEAE

A. Soriano 1984. 1/1. Arid regions of Argentina (from

La-Rioja to Santa-Cruz).

Halophytum.

A very specialized annual succulent herb that has

usually been included in the Chenopodiaceae and is

kept there even in Cronquist’s system (1981). However

from the Chenopodiaceae it differs markedly in the

ultrastructure of the sieve-element plastids, which pos-

sess a central globular crystal (Hunziker et al. 1974;

Behnke in Behnke and Mabry 1994). According to

Hunziker et al. (2000) Halophytaceae closely related

with Basellaceae, and perhaps also with the

Hectorellaceae and Portulacaceae, this idea also sup-

ported by the P-type of sieve-element plastids, the oral

structure and pollen morphology (Nowicke and Skvarla

1979) and ontogeny of stomata (De Fulvio, 1975).

14. CACTACEAE

A.L. de Jussieu 1789 (including Cereaceae De Candolle

et Sprengel 1821, Nopaleaceae Schmid et Curtman

1856, Opuntiaceae Desvaux 1817). About 100/1500 or

more. Arid and semiarid regions of America from

British Columbia and Alberta as far south as Patagonia,

concentrated in the dryer regions of Mexico, south-

western United States, and southern South America;

the only paleotropical member of the family is Rhip-

salis baccifera in Africa, Madagascar, Mascarenes,

Seychelles, and Sri Lanka.

14.1 PERESKIOIDEAE

Leaves large, supervolute, with lamina. Spines present,

but glochids absent. Tree-like, shrubby or scandent

plants. Ovary from superior to fully inferior. Funicles

ﬂ eshy in fruit. Seeds black and without an aril, unspe-

cialized, without complicated seed coat structure; they

resemble very much the seed coats of many

Portulacaceae (e.g., Claytonia) and Didiereaceae, e.g.,

Alluaudia) and support the ancestral position of the

family (Behnke and Barthlott 1983). – Pereskia

(including Rhodocactus).

14.2 MAIHUENIOIDEAE

Leaves small, terete, persistent; spines usually three.

Caespitose shrubs. Stems succulent, globose or short-

cylindric. Stomata parallelocytic. Flowers terminal,

solitary. Fruits obovoid or oblong, somewhat ﬂ eshy.

Seeds almost circular. – Maihuenia.

14.3 OPUNTIOIDEAE

Leaves and glochids present. Seeds covered by pale,

bony aril or winged, exhibiting particularly specialized

and unique type. This veriﬁ es the isolated position of

the subfamily, also indicated by its pollen morphology

(Behnke and Barthlott 1983). – Pereskiopsis, Quia-

bentia, Tacinga, Opuntia, Pterocactus.

14.4 CACTOIDEAE

Leaves absent or very minute, glochids absent. Seeds

black or brown, not enveloped by a bony aril. The

Cactoideae display a great and complex diversity of

seed coat characters that reﬂ ects their advanced posi-

tion within the family. – echinocereeae: Leptocereus,

Harrisia, Acanthocereus, Peniocereus, Echinocereus;

hylocereeae: Weberocereus, Hylocereus, Selenicereus,

Epiphyllum, Disocactus, Pseudorhipsalis; cereeae:

Cereus, Cipocereus, Brasilicereus, Piloso cereus,

Stephanocereus, Arrojadoa, Micranthocereus,

Coleocephalocereus, Melocactus; trichocereeae:

Cleistocactus, Leocereus, Haageocereus, Brachycereus,

140 Subclass IV. CARYOPHYLLIDAE

Espostoa, Espostoopsis, Facheiroa, Samaipaticereus,

Cleistocactus, Denmoza, Oreocereus, Matucana, Oroya,

Echinopsis, Arthrocereus, Rebutia, Mila, Gymnocaly-

cium, Discocactus; notocacteae: Corryocactus,

Austrocactus, Eulychnia, Copiapoa, Neowerdermannia,

Eriosyce, Neoporteria, Parodia, Blossfeldia, Frailea,

Uebelmannia; rhipsalideae: Lepismium, Rhipsalis,

Hatiora, Schlumbergera; browningieae: Calymman-

thium, Armatocereus, Jasminocereus, Neoraimondia,

Browningia, Stetsonia; pachycereeae: Bergerocactus,

Pachycereus, Carnegiea, Neobuxbaumia, Cephalo-

cereus, Stenocereus, Rathbunia, Polaskia, Escontria,

Myrtillocactus; cacteae: Echinocactus, Geohintonia,

Astrophytum, Sclerocactus, Pediocactus, Thelocactus,

Coryphantha, Neolloydia, Mammilloydia, Epithel-

antha, Ariocarpus, Lophophora, Obergonia, Strombo-

cactus, Aztekium, Pelecyphora, Ferocactus,

Leuchtenbergia, Stenocactus, Escobaria, Ortegocactus,

Mammillaria.

Morphological studies over the last few decades

(and likewise chemotaxonomic data) have conﬁ rmed

the close afﬁ nity of the Cactaceae with the

Phytolaccaceae, Portulacaceae, Didiereaceae, and

Aizoaceae. As early as 1862 Bentham and Hooker

included Cactaceae together with the Ficoideae

(Aizoaceae) in the order Ficoidales and thus correctly

showed their afﬁ nity. In Hauler’s and Wettstein’s sys-

tems the Cactaceae stand between Aizoaceae and

Portulacaceae. Numerous morphological data, accu-

mulated during the last decades, fully conﬁ rm the

closeness of Cactaceae to the Phytolaccaceae,

Aizoaceae, and Portulacaceae (see Chorinsky 1931;

Mauritzon 1934; Martin 1946; Buxbaum 1962;

Erdtman 1952; Barthlott and Hunt 1993). The relation-

ship of the Cactaceae with the other members of

Caryophyllales is supported also by phytochemical

data (Reznik 1955; Hegnauer 1964, 1989). The sub-

family Pereskioideae is in some respects an intermedi-

ate group between the apocarpous ancestors and more

advanced subfamilies of the Cactaceae.

15. DIDIEREACEAE

Radlkofer 1896 (including Portulacariaceae Doweld

2001). 7/16. South Africa (Portulacaria and Ceraria),

trop. Africa (Calyptrotheca), southern and south

western Madagascar.

15.1 DIDIEROIDEAE

Very spiny xerophytic shrubs or trees. Leaves alter-

nate, sometimes much reduced. Ovary (2)3–4-locular,

but usually only one locule is fertile. Fruit dry, indehis-

cent. – Decaryia, Alluaudiopsis, Alluaudia, Didierea.

15.2 PORTULACARIOIDEAE

Much-branched glabrous shrubs with waxy bark.

Leaves opposite, succulent. Flowers in cymose clus-

ters. Sepals persistent. Ovule solitary. Fruit a thin

walled nut with a single seed. – Portulacaria,

Ceraria.

15.3 CALYPTROTHECOIDEAE

Shrubs or small trees, sometimes climbing. Leaves

alternate, succulent with crinkled margins. Flowers in

panicles of cymes. Stamens up to 60. Ovules six.

Capsule circumscissile at the base, splitting upwards

into six valves. Seed solitary, arillate. – Calyptrotheca.

Have a common origin with the Cactaceae and

Portulacaceae and stand close to them. As early as

1908 and 1912 Hallier included Didiereaceae in the

Caryophyllales and put them near the Portulacaceae

s. 1. as their possible derivatives. According to

Erdtman (1948, 1952) pollen grains of the Didiereaceae

reveal many common features with the Nyctaginaceae

(Phaeoptilum and others) and stratiﬁ cation of the

exine resembles some pollen types of Cactaceae.

Close afﬁ nity with the Cactaceae is conﬁ rmed by the

resemblance in the structure of vegetative organs of

Didiereaceae and primitive members of the Cactaceae

as well as by the successful grafting of stems of

Didiereaceae to Pereskiopsis and Trichocereus (Rauh

and Reznik 1961). At the same time Didiereaceae

have much in common in ﬂ oral morphology with the

Portulacaceae (see Scholch 1963) and their epicuticu-

lar wax ultrastructure (irregular platelets and curled

ribbons) looks much the same as that of Aizoaceae

and Cactaceae (Engel and Barthlott 1988; Barthlott in

Behnke and Mabry 1994). In addition, embryo devel-

opment of the Didiereaceae is of the Chenopodium-

type, conﬁ rming their placement in the Caryophyllales

(Scholch 1963). The formation of betalains instead of

anthocyanins (Reznik 1955, 1975) and sieve-element

plastids with globular protein crystal (Behnke 1976b,

1978, 1982) as well as serological studies (Jensen

1965) also place the Didiereaceae in the order

Caryophyllales.

Superorder CARYOPHYLLANAE 141

16. MOLLUGINACEAE

Bartling 1825 (including Adenogrammaceae Nakai

1942, Glinaceae Dumortier 1829, Pharnaceaceae

Martynov 1820, Polpodaceae Nakai 1942, Telephiaceae

Martynov 1820). 12/120. Mainly tropical and subtrop-

ical regions, especially in Africa.

Corbichonia, Macarthuria, Psammotropha, Adeno-

gramma, Glischrothamnus, Mollugo, Glinus, Hypertelis,

Pharnaceum, Suessenguthiella, Coelan thum, Polpoda.

Related to the Phytolaccaceae, Aizoaceae, and

especially to the Sesuviaceae, differing mainly in

the absence of betalains. Also, they differ from the

Aizoaceae and Sesuviaceae in the presence of ﬂ a-

vonols and of C-glycosyl-ﬂ avonoids (viteksin and

isoviteksin) (Richardson 1978, 1981). The plastids

of Glinus include some starch grains (Behnke 1976b;

Behnke et al. 1983a, b), which is rare in the order. In

all other Molluginaceae the plastids have globular pro-

tein crystals.

According to Endress and Bittrich (1993), the inclu-

sion of the poorly known South African genus Polpoda

in the Molluginaceae is dubious. The stylodia are con-

nate at the base, a condition not found in the

Molluginaceae (Hofmann in Behnke and Mabry 1994).

17. LIMEACEAE

Shipunov ex Reveal 2005. 1/21. Southern Africa, to

Ethiopia, S. Asia.

Limeum.

Related to Molluginaceae, but differs in a lacking of

pigments and type of sieve-element plastids (Behnke

et al. 1983).

18. CARYOPHYLLACEAE

A.L. de Jussieu 1789 (including Alsinaceae Bartling

1825, Cerastiaceae Vest 1818, Corrigiolaceae

Dumortier 1829, Dianthaceae Vest 1818, Herniariaceae

Martynov 1820, Illecebraceae R. Brown 1810,

Lychnidaceae Löll 1843, Ortegaceae Martynov 1820,

Paronychiaceae A. L de Jussieu 1815, Polycarpaeaceae

Martens 1835, Scleranthaceae J. Presl et C. Presl 1820,

Silenaceae Bartling 1825, Spergulaceae Bartling 1825,

Stellariaceae Berchtold et J. Presl 1820, Telephiaceae

Martynov 1820). 104/ 2400. Mainly temperate regions

of the Northern Hemisphere with some species in the

Arctic, in the temperate regions of the Southern

Hemisphere, and on mountains in the tropics. The cen-

ter of distribution is in the Mediterranean region and

adjoining parts of Europe, and in western and Central

Asia. All the larger genera are found in the Northern

Hemisphere.

18.1 ILLECEBRIOIDEAE

Arnot 1832 (PARONYCHIOIDEAE A. Saint-Hilaire

1839). Stipules almost always present, mostly scari-

ous. Leaves alternate. Sepals free or slightly connate.

Petals mostly inconspicuous or absent. Pollen grains

3-colpate (rarely 3-colporate in Polycarpaea spp.) to

pantocolpate and pantoporate. – sperguleae:

Spergula, Spergularia; telephieae: Telephium; cor-

rigioleae: Corrigiola; xerotieae: Xerotia; poly-

carpaeae: Drymaria, Sanctambrosia, Polycarpon,

Polycarpaea, Ortegia, Loeﬂ ingia, Haya, Krauseola,

Polytepalum, Stipulicida, Cerdia, Microphyes, Pirinia;

paronychieae: Cometes, Dicheranthus, Pteranthus,

Sphaero coma, Sclerocephalus, Lochia, Paronychia,

Herniaria, Philippiella, Chaetonychia, Achyronychia,

Illecebrum, Cardionema, Scopulophila, Pollichia.

18.2 MINUARTIOIDEAE (ALSINOIDEAE)

Stipules absent. Leaves opposite. Sepals free or rarely

connate at the base. Petals present or rarely absent.

Fruits mainly capsules, rarely indehiscent. – alsineae:

Arenaria, Dichodon, Eremogone, Thylaco spermum,

Moehringia, Brachystemma, Thurya, Bufonia,

Lepyrodiclis, Cerastium, Stellaria, Mesoste mma,

Tytthostemma, Fimbripetallum, Pseudostellaria,

Moenchia, Holosteum, Myosoton, Minuartia, Queria,

Wilhelmsia, Honkenya, Sagina, Colobanthus, Alsiniden-

dron, Schiedea, Reicheella, Plettkea, Pycnophyllopsis;

pycnophylleae: Pycnophyllum; SCLERANTHEAE:

Scleranthus, Pentastemonodiscus; Sgeocarpeae:

Geocarpon; HABROSIEAE: Habrosia.

18.3 CARYOPHYLLOIDEAE

Stipules absent. Sepals connate. Petals present, often

clawed. – caryophylleae: Acanthophyllum,

Kuhitangia, Allochrusa, Scleranthopsis, Ochotono-

phila, Diaphanoptera, Gypsophila, Psammophiliella,

Dichoglottis, Vaccaria, Pleioneura, Saponaria,

Spanizium, Bolbosaponaria, Pseudosaponaria,