Takhtajan Armen. Flowering Plants

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152 Subclass IV. CARYOPHYLLIDAE

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Order 43. PHYSENALES

Small trees or shrubs. Vessels with simple perforations

with slightly inclined end walls; lateral pitting alternate.

Rays homogenous or heterogenous. Fibers with simple

to bordered pits. Axial parenchyma aliform to conﬂ u-

ent and distinctly unilateral, paratracheal. Sieve-element

plastids of P-type. Nodes unilacunar. Leaves alternate,

entire, estipulate. Stomata anomocytic. Flowers in

racemes or panicles, or solitary, unisexual or bisexual.

Sepals 5–9, slightly unequal, imbricate, persistent, each

sepal vascularized by a single trace; petals 5 or absent.

Stamens 8–25 (usually 10–14), essentially in one

cycle; ﬁ laments free, short, slightly connate at the

base; anthers basiﬁ xed or dorsiﬁ xed, tetrasporangiate,

latrorse, opening longitudinally, with endothecal-like

connective hypodermis. Pollen grains in monads,

2–3-colpate or sometimes 6-colpate. Tectum traversed

Superorder CARYOPHYLLANAE 153

by numerous microchannels, appearing on the surface

as microperforations; ornamentation is in the form of

small spines or spinules; endexine lamellate in both

aperture and nonaperturate regions. Gynoecium of 2–3

united carpels. Ovary superior, mainly 2–3-locular,

with 4–5 pendulous, campylotropous, epitropous,

bitegmic (with the inner integument longer than the

outer), crassinucellate ovules. Fruits dry, indehiscent.

Seeds large, with straight or curved embryo, without

endosperm. Plants producing betalain.

The sister-group relationship of Physena and

Asteropeia was demonstrated by Morton et al. (1997),

and the sister relationship of this clade to the remain-

ing core Caryophyllales has been conﬁ rmed in several

studies (e.g., Morton et al. 1997; Soltis et al. 2000,

2006; Guénoud et al. 2002).

Key to Families

1 Sepals enlarged and winglike in fruit. Fibers with

small bordered pits. Rays homogenous. Small to

tall, evergreen trees and scrambling shrubs. Leaves

simple, shortly petiolate. Flowers bisexual, in axil-

lary or terminal panicles. Petals ﬁ ve, adnate to the

calyx, imbricate. Stamens (9)10–15; ﬁ laments

dilated and connate at the base; anthers small and

versatile, short- dorsiﬁ xed. Pollen grains 3-colpate

or 6-colpate, rarely 2-colpate, ﬁ nely spinulate.

Gynoecium of 3(2) carpels; style usually with 3 sty-

lar branches or with 3-lobed stigma. Ovary usually

incompletely 3-locular or rarely 2-locular, with 2 or

less often 4–5 (rarely 6) pendulous, epitropous ovules

in each locule. Fruits thick-walled, dry, surrounded

by persistent stamens and strongly accrescent and

winglike sepals. Seeds reniform to hippocratiform

with radicle close to large hilum; embryo curved,

sometimes sigmoid; cotyledons large and thin, spi-

rally coiled……………… 1. asteropeiaceae.

1 Sepals not enlarged and winglike in fruit. Shrub or

tree. Fibers with simple or minutely bordered pits.

Rays homogeneous typically with square and/or

upright cells, or heterogeneous with more than four

rows of upright and/or square cells. Leaves coria-

ceous, simple, with pinnate venation, and arch-

shaped petiole bundle. Flowers on long pedicels in

5–10-ﬂ owered axillary racemes (Physena madagas-

cariensis) or sessile or nearly sessile and solitary, or

in 2–3-ﬂ owered axillary clusters (P. sessiliﬂ ora), uni-

sexual (dioecious), actinomorphic, apetalous, ane-

mophilous. Stamens (8)10-14(-25); ﬁ laments short,

slightly connate at the base; anthers long, linear, bas-

iﬁ xed. Pollen grains 3-colpate or sometimes with

4–5 short meriodinal colpi; ornamentation is in the

form of small spines or spinules. Male ﬂ owers with a

pistillodium. Gymoecium of two carpels. Stylodia

very long, ﬁ liform, arising from the depression at the

top of the ovary, connate only at the very base, papil-

late surface nearly along their whole length. Ovary

2-locular at the base and apex and locular at the

midlevel, with four ovules (two ovules per locule) on

axile placentas, each with a long funicle directed

upward. Ovules campylotropous. Fruits dry, indehis-

cent, 1-seeded, with coriaceous and somewhat

inﬂ ated pericarp that is separated from the seeds.

Seeds large, with straight embryo consisting of two

massive cotyledons of unequal size and minute,

punctiform radicle. . . . . . . . . . . . . . 2. physena ceae

1. ASTEROPEIACEAE

Takhtajan ex Reveal et Hoogland 1990. 1/8. Madagascar.

Asteropeia.

2. PHYSENACEAE

Takhtajan 1985. 1/2. Madagascar.

Physena.

According to Miller and Dickison (1992), the wood

of Physena so much resembles that of Asteropeia that

“solely on wood anatomy, these two genera could be

placed in the same family.” It is all the more important,

then, that many years ago Erdtman (1952) noticed simi-

larities between pollen grains of Physena and Asteropeia.

Morton, Karol, and Chase (1997) also mentioned many

similarities between Asteropeia and Physena, and con-

sider them other sister group to the Caryophyllales.

Bibliography

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Adansonia, ser. 2, 8: 355–357.

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A case study in comparative wood anatomy. Brittonia 58:

301–313.

Dickison WC and RB Miller. 1993. Morphology and anatomy of

the Malagasy genus Physena (Physenaceae), with a discussion

of the relationships of the genus. Adansonia 1–4: 85–106.

154 Subclass IV. CARYOPHYLLIDAE

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ics from Madagascar. Am. J. Bot. 79(6): 83 (Abstract.)

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relationships of two anomalous dicot genera Physena and

Asteropeia: evidence rbcL plastid DNA sequences. Bot. Rev.

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sclereids in Asteropeia Thou. Curr. Sci. 53: 45–48.

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of Madagascar, IV. A synoptic revision of Asteropeia

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Superorder POLYGONANAE

Order 44. POLYGONALES

Perennial or annual herbs, shrubs, lianas, or trees, often

with conspicuous swollen nodes. Woody species often

with anomalous secondary growth; silica bodies and

calcium oxalate crystals present in wood cells. Vessels

with simple perforations; lateral pitting alternate; ves-

tured pits mostly absent (except Brunnichia,

Muehlenbeckia, Triplaris, and Coccoloba – Carlquist

2003). Fibers with simple pits, mostly septate. Axial

parenchyma scanty paratracheal or vasicentric. Rays

from homogeneous to heterogeneous. Sieve-element

plastids of S-type or (Triplaris, Ruprechtia, Coccoloba)

Pfs-type (Behnke 1999). Nodes multilacunar or less

often trilacunar. Leaves alternate, rarely opposite or

verticillate, entire or rarely pinnatiﬁ d or palmately cleft,

membranous to ﬂ eshy or leathery, sometimes articu-

lated at the base; stipules interpetiolar, connate into a

usually membranous or scarious sheath (ochrea) around

the stem and completely or partly covering the nodes,

reduced to a row of hairs or wanting in Eriogonoideae

and in a few members of Polygonoideae. Stomata com-

monly anomocytic, sometimes anisocytic, diacytic, or

paracytic. Flowers small, seldom solitary, mostly in

various types of inﬂ orescences, bisexual or less often

unisexual and dioecious, actinomorphic, 3-merous or

5-merous, rarely 2-merous, apetalous. Sepals (2-)5 or 6,

rarely 9 (Rheum), imbricate or quincuncial in bud, green

and herbaceous to often colored and petaloid, more or

less connate at the base or forming a short or long ﬂ oral

tube, persistent and sometimes enlarging in fruit.

Stamens (3)5–8(9); ﬁ laments ﬁ liform, free or basally

connate; anthers tetrasporangiate, 2-locular, versatile or

basiﬁ xed, opening longitudinally. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains 3-celled

or rarely 2-celled, mostly 3-colporate, rarely panto-

hexocolporate (Hong et al. 2005). A nectary disc pres-

ent at the base of the ovary, which is annular as in

Caryophyllaceae, or nectaries are placed between the

bases of the ﬁ laments. Gynoecium of three or less often

two, rarely four, carpels, with free or more or less con-

nate stylodia; ovary superior, subtended by an annular

(often lobed) glandular disc, unilocular, sometimes

with vestigial partitions at the base. Ovule solitary,

basal on more or less well-expressed stalk representing

a reduced free-central placenta, orthotropous or rarely

anatropous, bitegmic or sometimes more or less uniteg-

mic through connation (Corner 1976), crassinucellate.

Female gametophyte of Polygonum-type. Endosperm

nuclear. Fruits nutlike, very often trigonous, sometimes

closely subtended by the persistent calyx or enclosed in

a ﬂ eshy ﬂ oral tube. Seeds of medium size, straight; seed

coat formed by the outer integument (testal); embryo

straight or more often curved, well-differentiated, sur-

rounded by a copious, starchy, and oily, horny, some-

times ruminate endosperm with solitary starch grains;

perisperm essentially lacking. Commonly tannin-

iferous, producing anthocyanins but not betalains,

sulphated ﬂ avonols, ellagic acid, n = 7–13.

Polygonales are related to the Caryophyllales and

probably derived from some of their archaic members

with 3-colporate pollen grains, apetalous 3-merous

ﬂ owers, and stipulate leaves. Bessey (1915) included

Polygonaceae in the Caryophyllales between

Chenopodiaceae and Nyctaginaceae, while Hallier

(1908, 1912) derived the family from Portulacaceae s.

1. (including Basellaceae) and Chenopodiaceae. There

are many features in common between the

Polygonaceae and Portulacaceae, especially in the

morphology of the gynoecium, which in both families

usually consists of three carpels and is of lysicarpous

type. In the seeds of Caryophyllaceae there are ves-

tiges of nucellar tissue with very small starch grains,

which are characteristic of the perisperm of

Caryophyllales. The amino acid sequence in members

of the Polygonaceae and Caryophyllales also conﬁ rms

their afﬁ nity (Boulter 1973). But at the same time the

Polygonales (as well as the next order Plumbaginales)

Superorder POLYGONANAE 155

differ from the Caryophyllales in their S-type sieve-

element plastids and from most of them also in their

lack of betalains.

1. POLYGONACEAE

A. L. de Jussieu 1789 (including Calligonaceae

Chalkuziev 1985, Eriogonaceae G. Don 1839,

Persicariaceae Martynov 1820, Rumicaceae Martynov

1820). 54/c.1100. Very widely distributed, but chieﬂ y

in northern temperate regions; quite a few are tropical,

including Antigonon, Coccoloba, and Muehlenbeckla.

1.1 ERIOGONOIDEAE

Leaves without well-deﬁ ned stipules (ochrea lacking).

Branching often sympodial. Inﬂ orescences cymose.

Flowers always 3-merous. Anthraquinones probably

lacking. — eriogoneae: Eriogonum, Oxytheca,

Dedeckera, Gilmania, Stenogonum, Goodmania,

Nemacaulis, Dodecahema, Aristocapsa, Chorizanthe,

Mucronea, Systenotheca, Centrostegia, Hollisteria,

Lastarriaea; pterostegieae: Pterostegia, Harfordia.

1.2 POLYGONOIDEAE.

Leaves with stipular sheaths (ochrea more or less

developed). Branching monopodial. Inﬂ orescences

racemose with cymose partial inﬂ orescences. Flowers

3-merous, 2-merous, or 5-merous. Anthraquinones

often present. – rumicieae: Rheum, Rumex, Oxyria,

Emex; persicarieae: Persicaria, Antenoron, Bistorta,

Aconogonon, Koenigia, Fagopyrum; polygoneae:

Polygonum, Cephalolophilon, Truellum, Knorringia,

Chylocalyx Polygonella, Reynoutria, Fallopia,

Oxygonum; atraphaxideae: Atraphaxis, Pteropyrum,

Calligonum, Physopyrum, Para pteropyrum; triplar-

ideae: Gymnopodium, Lepto gonum, Ruprechtia, ?

Enneatypus, Triplaris, Symmeria; coccolobeae:

Muehlenbeckia, Homalocladium, Coccoloba,

Antigonon, Brunnichia, Afrobrunnichia, Podopterus,

Neomillspaughia.

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Order 45. PLUMBAGINALES

Perennial or sometimes annual herbs, subshrubs, or

low shrubs, rarely lianas. Secondary growth in some

genera (including Acantholimon, Aegialitis, and

Limoniastrum) anomalous. Vessels with simple perfo-

rations; lateral pitting alternate. Fibers with simple

pits. Rays homogeneous. Axial parenchyma scanty

paratracheal. Sieve-element plastids of S-type. Nodes

trilacunar. Leaves alternate, entire, usually estipulate,

only rarely with well-developed stipules, often with

scattered chalk-glands exuding water and salts, and

often also raised mucilage glands in leaf axils and on

the surface. Stomata anomocytic, or anisocytic, or

paracytic. Flowers in cymose heads or panicles or in

racemes or spikes, bisexual, actinomorphic, 5-merous,

with both sepals and petals, bracteate (the bracts some-

times forming on involucre), with 2(1) bracteoles,

often heterostylic. Sepals connate to form a plicate,

often conspicuously 5-ribbed or 10-ribbed tube, the

lobes mostly dry and membranous or scarious, often

showy, persistent. Petals connate at the very base or

less often into a tube, often persistent; the petals or

corolla lobes contorted or imbricate. Stamens ﬁ ve,

opposite the petals or corolla lobes, or partly adnate to

the tube; anthers dorsiﬁ xed or (Aegialitis) basiﬁ xed,

tetrasporangiate, introrse, 2-locular, opening longitu-

dinally. Tapetum secretory. Microsporogenesis simul-

taneous. Pollen grains 2-celled or more often 3-celled,

tectate-columellate, 3-colpate or rarely 4–5-colpate,

very rarely pantocolpate, usually dimorphic in the

Staticoideae. Gynoecium of ﬁ ve united carpels; stylo-

dia more or less distinct (Staticoideae) or connate into

an apically lobed style (Plumbaginoideae); stigmas

papillate; ovary superior, unilocular, usually 5-lobed

or -ribbed, with solitary basal ovule on elongate, slen-

der funicle that almost encircles it. Ovules pendulous,

anatropous, sometimes circinotropous, bitegmic,

crassinucellate. Female gametophyte tetrasporic, 4–8-

nucleate, of Fritillaria-, Adoxa-, Plumbagella-,

Plumbago- and Penaea-type. Endosperm nuclear.

Fruits mostly more or less enclosed within the persis-

tent calyx, dry, indehiscent or less often tardily cir-

cumscissally dehiscent or splitting upward into valves.

Seeds small, winged; seed coat formed by both integu-

ments and consists of exotesta and endotegmen (Vovk

1991). Embryo rather large, green, straight, well-

differentiated; endosperm starchy, more or less

copious or scanty, sometimes (as in Aegialitis and

Psylliostachys) absent. Perisperm lacking. Plants often

salt tolerant; producing various 0-methylated ﬂ avonols

and anthocyanidins, glycine betaine, choline-O-sul-

phate, plumbagin, kaempferol, quercetin, and myrice-

tin, but no betalains, generally tanniniferous. n = 6–9.

Agardh (1858) put Plumbaginaceae close to the

Nyctaginaceae and Hallier (1901, 1903, 1908, 1912)

derived Plumbaginaceae directly from Portulacaceae

s. 1. (including Basellaceae). With both the

Portulacaceae and Basellaceae they share the important

Superorder POLYGONANAE 157

feature of a lysicarpous gynoecium, which in

Plumbaginaceae and Basellaceae is characterized by a

reduced free central column. Friedrich (1956) brought

together many gross morphological, anatomical,

palynological, embryological, and cytological data

that conﬁ rm the afﬁ nity of the Plumbaginaceae with

the families of “Centrospermae.” He derived

Plumbaginaceae directly from the Phytolaccaceae,

which seems unlikely.

The Plumbaginaceae differ from the Caryophyllales

in so many features that they deserve the status of a

separate order. They differ from them by S-type plas-

tids, pollen grain morphology, generally sympetalous

corolla, the absence of perisperm and presence of

endosperm, and straight embryo. The rbcL sequence

data indicate that the Polygonales and Plumbaginales

should be regarded as sister groups perhaps evolved

parallel to, rather than directly related to the

Caryophyllales (Giannasi et al. 1992).

1. PLUMBAGINACEAE

A. L. de Jussieu 1789 (including Aegialitidaceae

Linczevsky 1968, Armeriaceae Horaninow 1834,

Limoniaceae Seringe 1851, Staticaceae Cassel 1817).

29/750–850. Almost cosmopolitan but especially well

represented in dry and saline habitats, particularly in

the Mediterranean and Irano-Turanian regions.

1.1 PLUMBAGINOIDEAE

Flowers in racemes or spikes. Sepals herbaceous, more

or less glandular. Corolla sympetalous. Stamens mostly

free or partly connate to the corolla. Pollen grains with

well-deﬁ ned foot-layer, highly irregular columellae, and

a complete tectum. Stylodia connate into an apically

lobed style. Naphtoquinone plumbagin, glycine betaines,

5-O-methylated ﬂ avonols always present, n = 6–7. –

Plumbago, Plumbagella, Dyerophytum, Ceratostigma.

1.2 STATICOIDEAE

Flowers in panicles or heads with monochasial partial

inﬂ orescences or rarely (Aegialltis) in leafy, often fork-

branched racemes. Sepals scarious. Petals connate only at

the very base, rarely (Psylliostachys) corolla sympeta-

lous. Filaments basally adnate to the petals. Pollen grains

with a foot-layer supporting straight, regular columellae,

which are distally fused into an incomplete rectum.

Stylodia free or rarely (Limoniastrum) connate up to the

middle. Present betaalanine betaines, glycine betaines

rare, naphtoquinone plumbagin lacking, n = 8, 9. –

Aegialitis, Acantholimon, Neogontscharovia,

Gladiolimon, Ghaznianthus, Dictyolimon, Cephalorhizum,

Bamiania, Popoviolimon, Chaetolimon, Vassilczenkoa,

Limoniopsis, Ikonnikovia, Goniolimon, Bukiniczia,

Afrolimon, Bakerolimon, Muellerolimon, Limoniastrum

(including Bubania), Limonium, Myriolimon,

Saharanthus, Eremolimon, Armeria, Psylliostachys.

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207–219.

Baker HG. 1953. Dimorphism and monomorphism in the

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niﬁ cance. Belg. J. Bot. 125: 151–153.

Dahlgren KV0. 1916. Zytologische und embryologische Studien

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158 Subclass IV. CARYOPHYLLIDAE

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Roth I. 1962a. Histogenese und morphologische Deutung der

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19–40.

Roth I. 1962b. Histogenese und morphologische Deutung der

Kronblatter von Armeria. Port. Acta Biol, ser. A., 6:

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Schoute JC. 1935. Observations on the inﬂ orescence in the fam-

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406–424.

Vovk AG. 1991. Plumbaginaceae. In: A Takhtajan, ed.

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Leningrad (in Russian).

Weberling F. 1956. Weitere Untersuchungen zur Morphologie

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Plumbaginaceae. Beitr. Biol. Pﬂ . 33: 17–32.

Wolf GR, S Xu, GW Patterson, and TA Salt. 1989. Polygonales

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Caryophyllidae. Phytochemistry 28: 143–145.

Order 46. TAMARICALES

Trees, shrubs, subshrubs or less often perennial herbs,

mostly halophytic or xerophytic with storied wood.

Vessels with simple perforations; lateral pitting alter-

nate. Fibers mostly with few, simple pits. Rays in

Tamaricaceae very broad and high, heterogeneous, but

in Frankeniaceae deﬁ nite medularry rays are lacking.

Axial parenchyma in Tamaricaceae scanty- paratracheal

to vasicentric. Sieve-element plastids of S-type. Leaves

alternate or opposite simple, entire, mostly small, com-

monly with embedded, multicellular, salt-excreting

glands, estipulate. Stomata usually anomocytic. Flowers

mostly small or very small, solitary or more often in

various kinds of inﬂ orescences, usually bisexual, acti-

nomorphic. Sepals 4–7, free or less often more or less

connate (in Frankeniaceae united into a tube with lobes),

imbricate or valvate. Petals as many as the sepals, free,

imbricate. Nectary disc present or wanting. Stamens as

many as petals, free or more or less connate; anthers

tetrasporangiate, opening longitudinally. Tapetum secre-

tory. Microsporogenesis simultaneous. Pollen grains

2-celled or 3-celled, (2)3(4)-colpate, rarely rugate.

Gynoecium of 2–5 (mostly 3–4) united carpels with sty-

lodia free or only basally connate. Ovary superior,

1-locular, with (1)2-many ovules on each parietal pla-

centa. Ovules anatropous, bitegmic, more or less tenui-

nucellate (Tamaricaceae) or thinly crassinucellate

(Frankeniaceae – Fedotova 1992). Female gametophyte

of Polygonum-type or tetrasporic of various types

(Kamelina 1983). Endosperm nuclear, or rarely (Tamarix

tetrandra) cellular, without haustoria or with micropylar

haustorium (Frankeniaceae). Fruits loculicidal capsules.

Seeds small, covered with long uni cellular hairs or with

a chalazal hair tuft, or seeds uniformly papillate; seed

coat formed by the outer integument (exotestal); embryo

large, straight; endosperm copious and starchy, scanty

and starchy or lacking. Commonly producing bisul-

phated ﬂ avonols, but not cyanogenic.

According to molecular data Tamaricales are clos-

est to the Plumbaginales, which does not contradict

morphological data.

Key to Families

1 Leaves alternate, not ericoid or revolute- margined,

often scaly or subulate, commonly with external

but embedded, multicellular salt glands, simple,

estipulate. Stomata anomocytic or paracytic.

Flowers small to medium size, in racemes or in

spike, or in panicles, or solitary, bisexual or rarely

unisexual (dioecious). Sepals 4–5(6), free or less

often basally connate, imbricate. Petals 4–5, alter-

nate with sepals with two scalelike appendages

(Reaumurieae) or without appendages, free or

slightly connate, imbricate, persistent or caduceus.

Stamens 4–6, or 8–12 (often twice as many as the

petals), or 15–100 (often more or less numerous),

seated on a ﬂ eshy nectary disc, free or basally con-

nate; anthers 2-locular, dorsiﬁ xed, extrorse,

latrorse, or introrse, opening by longitudinal slits.

Pollen grains 2- or 3-celled. Gynoecium of 3–4,

rarely 2 or 5 united carpels; ovules ascending.

Female gametophyte tetrasporic, mostly of

Fritillaria-type. Seeds covered with long unicellular

Superorder POLYGONANAE 159

hairs; exotestal cells periclinally elongated and

thick- walled, endotestal cells thin-walled, crystal-

liferous; endosperm scanty or lacking. Trees,

shrubs or subshrubs. Gypsum crystals present.

Contain kaemp ferol, quercetin, or tamatixin, alka-

loids present or absent. n = 11 (Reaumurieae) or

12 (Tamariceae). . . . . . . . . . . . 1. tamaricaceae

1 Leaves opposite, often ericoid and revolute- margined,

commonly punctuate with sunken, salt- excreting

glands. Medullary rays wanting. Flowers solitary and

sessile in the forks of the branches or forming a dense

leafy cyme. Sepals 4–7, connate into a tube with

short, induplicate-valvate lobes. Petals 4–7, distinct,

imbricate, clawed, usually with a scale-lake append-

age inside. Nectary disc wanting. Stamens 4–7, or up

to 24, most often 6 in 2 cycles, free or shortly connate

at the base with ﬁ liform or ﬂ attened ﬁ laments; anthers

versatile, opening length-wise. Pollen grains 3-celled,

(2)3(4)-colpate. Gynoecium of (1-)3(-4) carpels

united to form a 1-locular ovary; style slender end

elongate, usually with distinct stylodia; ovules (1)2–

6(-many) in each carpel. Female gametophyte of

Polygonum-type. Seeds uniformly papillate, exot-

estal cells large, papilla with terminal nail-like thick-

enings, endotestal cells thin-walled, endotegmen

with thick cuticle, tanniniferous; embryo straight,

endosperm copious. Halophytic shrubs, subshrubs or

herbs. Contain tannins, proanthocyanidins, ellagic

acid, ﬂ avonols. n = 10, 15. . . . . . 2. frankeniaceae

1. TAMARICACEAE

Link 1821 (including Reaumuriaceae Ehrenberg ex

Lindley 1830). 5/90. Africa and Eurasia, mainly

Mediterranean and Irano-Turanian regions.

reaumurieae: Reaumuria, Hololachna; tama-

riceae: Tamarix, Myricaria, Myrtama.

2. FRANKENIACEAE

Gray 1821 1/90. Distributed throughout warmer dry

regions, especially on maritime shores of America,

Africa, Eurasia, and Australia, but best developed in

the Mediterranean region and western Asia.

Frankenia (including ? Hypericopsis, Beatsonia,

Anthobryum, Niederleinia).

Related to the Tamaricaceae and especially to the

Reaumurieae, which is conﬁ rmed also by a great simi-

larity in the ultrastructure of the salt glands (Campbell

and Thomson 1976), as well as secondary chemistry,

and often petaline scales, and also the molecular analy-

sis (Gaskin et al. 2004).

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Superorder NEPENTHANAE

Order 47. NEPENTHALES

Herbaceous or sometimes more or less woody plants

with normal or anomalous secondary growth, erect,

prostrate, or often twining or climbing, aquatic, terres-

trial or epiphytic, glabrous or hairy, cornivorous or not.

Vessels with simple perforation, true tracheids present.

Rays homogeneous or heterogeneous. Axial paren-

chyma paratracheal (vasicentric) and apotracheal,

banded and diffuse. Nodes unilacunar, trilacunar to

multilacunar (Nepenthaceae). Leaves alternate, simple,

estipulate or with small stipules, Stomata anomocytic,

tetracytic, actinocytic or encyclocytic. Flowers usually

small, terminal or axillary, in racemes, panicles, spikes,

or rarely (Aldrovanda) solitary, bisexual or rarely

(Nepenthaceae) unisexual, actinomorphic. Sepals usu-

ally 4–5 rarely 3, free or less often basally connate,

imbricate or rarely valvate, sometimes persistent. Petals

4–5, free or shortly connate, or cohering, imbricate or

contorted. Stamens 2–8(-25), rarely 4–6; ﬁ laments free

or united; anthers 2-locular, basiﬁ xed or dorsiﬁ xed,

extrorse, introrse or latrorse, tetrasporangiate, opening

longitudinally. Tapetum secretory. Microsporogenesis

simultaneous. Pollen grains 2–3-celled, in tetrads or

rarely (Drosophyllum) in monads, 3-colpate or 3–4-

colporate. Gynoecium of 2 or 5, very rarely 3–4 or 6

united carpels; stylodia more or less free or united;

stigma capitate, hippocrepiform or puntoﬁ d, or discoid,

on a very short style, or sessile; ovary superior to infe-

rior, 1-locular, or rarely 3–4-locular; ovules solitary

(Ancistrocladus) or numerous, anatropous or hemitrop-

ous, bitegmic, mostly crassinucellate. Female gameto-

phyte of Polygonum-type. Endosperm nuclear or cellular

(Ancistrocladus). Fruits loculicidal or valvular capsules,

or indehiscent (Aldrovanda and Ancistrocladus). Seeds

small to large, sometimes numerous (100–500), often

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

either end), or wingless; seed coat formed by both the

integuments, but mainly exotestal. Embryo well-differ-

entiated, straight, cylindrical or discoid-obconix, sur-

rounded by starchy, proteinaceous and oily endosperm,

sometimes endosperm absent. Produced proanthocya-

nidins, ﬂ avonols, acetogenic napthoquinone plumbagin,

and isoquinoline alkaloids.

Probably related to the some families of the

Polygonanae. “Possible synapomorphies for the noncore

Superorder NEPENTHANAE 161

Caryophyllales are scattered secretory cells containing

plumbagin, naphthaquinone (which has been lost in

several clades), an indumentum of stalked, gland-headed

hairs, basal placentation (with shifts to parietal in some

Droseraceae and axile in Nepentaceae), and starchy

endosperm” (Judd et al. 2002, Soltis et al. 2006)

Key to Families

1 Ovary mostly superior.

2 Flowers bisexual; petals imbricate or contorted.

3 Plants with normal secondary growth.

4 Perennial or annual carnivorous herbs,

commonly growing in bogs and other

waterlogged soils containing little or no

available nitrogen, rarely (Aldrovanda)

submerged aquatics, sometimes with

corms or root tubers; glandular hairs irri-

table when present. Nodes unilacunar with

1 trace. Leaves spiral or rarely whorled

(Aldrovanda), often circinate in bud, usu-

ally in basal rosettes, mostly with intra-

petiolar stipules; the blade either modiﬁ ed

into an active trap that shuts when touched

(Dionaea, Aldrovanda) or provided with

sticky tentacles and sessile glands that

entrap and digest insects and other small

animals (Drosera). Stomata anomocytic,

tetracytic or actinocytic. Flowers in cymose

inﬂ orescences or (Aldrovanda) solitary,

bisexual, actinomorphic. Sepals (4)5(-8),

more or less connate at the base, imbricate,

persistent. Petals as many as the sepals,

free, imbricate or contorted. Stamens

(4)5(10–20), free or (Dionaea) basally

connate; anthers 2-locular, dorsiﬁ xed or

basiﬁ xed, extrorse, tetrasporangiate, open-

ing longitudinally; anther wall initially

with one middle layer. Pollen grains in tet-

rads, 3-colpate to more often periporate.

Gynoecium of three or ﬁ ve united carpels,

with more or less free or (Dionaea) united

stylodia that have terminal stigma. Ovary

superior, or sometimes nearly half-inferior,

1-locular, with three parietal placentas

(Dionaea) or with essentially basal pla-

centas. Ovules anatropous, bitegmic,

crassinucellate or tenuinucellate in some

Drosera species. Endosperm nuclear.

Fruits loculicidal or valvular capsules, or

rarely indehiscent (Aldrovandra). Seeds

usually minute, mostly fusiform; seed coat

usually formed by both the integuments;

every epidermal layer of the seed coat is

potentially specialized, the exotesta as a

palisade, the endotesta as thick-walled

crystal-cells, and the endotegmen as scle-

roic cuboid cells; embryo short, straight,

embedded in the copious, oily endosperm

rich in starch. Produced proanthocyanidins

(cyanidin and delphinidin), ﬂ avonols

(including myricetin, kaempferol and

quer cetin), acetogenic napthoquinones

plumbagin, and ellagic acid (Drosera, 2

sp.); n = 5–24. . . . . . . . . 5. droseraceae

4 Small insectivorous tap-rooted subshrubs,

sometimes branched; axes and leaves cov-

ered with rows of stalked mucilage, red ten-

tacles and irregularly distributed sessile

digesting glands. Leaves spiral, crowded,

narrow-linear, reversely circinate in bud,

estipulate. Inﬂ orescence terminal, few-ﬂ ow-

ered, thyrso- paniculate. Flowers large, short-

lived, remain open only a single day, bisexual,

actinomorphic, 5-merous. Sepals imbricate,

fused at base, glandular. Petals free, con-

torted, deciduous. Stamens ten, free; anthers

extrorse. Pollen grains pantoporate, in

monads. Gynoecium of ﬁ ve united carpels;

stylodia with capitate stigmas; ovary 1-locu-

lar; ovules numerous, basal, anatropous,

with long funicles attached to a basal, coni-

cal placenta. Fruits loculicidal, coriaceous

capsules, opening in upper half. Seeds

numerous, seed coat with endotestal crystal

layer and sclerotised endotegmen. Embryo

small, endosperm copious, ﬂ eshy and

starchy. Produced proanthocyanins, the

naphtoquinone plumbagin, ﬂ avone luteolin,

and inulin; n = 6. . . . .4. drosophyllaceae

3 Plants with anomalous secondary growth.

Insectivorous (Triphyophyllum), soft-wooded

lianas, climbing by pairs of hooks or tendrils

representing the forked tips of the leaf mid-

ribs, heterophyllous (Triphyophyllum, which

in addition to the hook-tipped climbing leaves

has ‘normal’ leaves without hooks, and others

which are wholly reduced to the midrib, cir-

cinnate and beset with numerous stalked and