Takhtajan Armen. Flowering Plants

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716 Subclass IV. COMMELINIDAE

Classiﬁ cation after T.M. Evans, R. B. Faden, M. G.

Simpson, and K. J. Sytsma (2000).

2.1 CARTONEMATOIDEAE

Stem collenchyma absent. Glandular microhairs lack-

ing. Raphide canals lacking or only next to the veins.

Flowers actinomorphic – cartonemateae: Cartonema;

triceratelleae: Triceratella.

2.2 COMMELINOIDEAE

Stem collenchyma present. Glandular microhairs

nearly always present. Raphide canals present and

mostly between the veins of the lamina. Flowers acti-

nomorphic to zygomorphic — tradescantieae:

Palisota, Streptolirion, Spatholirion, Aetheolirion,

Cyanotis (including Amischophacelus), Belosynapsis,

Coleotrype, Porandra, Amischotolype, Dichorisandra,

Siderasis, Geogenanthus, Cochliostema, Plowma-

nianthus, Thyrsanthemum, Gibasoides, Tinantia,

Elasis, Matudanthus, Weldenia, Gibasis, Tradescantia,

Callisia, Tripogandra, Sauvallea; commelineae:

Stanﬁ eldiella, Floscopa, Buforrestia, Murdannia,

Anthericopsis, Tricarpelema, Pseudoparis, Polyspatha,

Dictyospermum, Pollia, Aneilema, Rhopalephora,

Commelina.

3. PHILYDRACEAE

Link 1821. 4/5–6. Eastern and Southeast Asia, Guam,

Andaman Islands, New Guinea, Australia (all 4 genera

and 5 species occur in Australia).

Helmholtzia, Orthothylax, Philydrum, Philydrella.

Philydraceae are related to the Pontederiaceae. The

common features of the Philydraceae and Pontederi-

aceae include the ovary position, embryology (embryo

and endosperm in the ripe seed), vegetative anatomy

( paracytic stomata, styloid crystal idioblasts, etc.) and

similar proanthocyanins (see Hamann 1966, 1998).

4. PONTEDERIACEAE

C. Kunth 1816 (including Heterantheraceae J. Agardh

1858, Unisemataceae Raﬁ nesque 1837). 9/33. Wide-

spread in tropics, with a few species extending into

warm-temperate areas; the centre of diversity is tropical

and subtropical America.

eichhornieae: Eichhornia, Monochoria; ponted-

erieae: Pontederia, Reussia; heteranthereeae:

Heter anthera, Zosterella, Hydrothrix, Scholleropsis,

Eurystemon.

Related to the Philydraceae, but even closer to the

Haemodoraceae (see Dahlgren and Rasmussen 1983

and Simpson 1987). According to Simpson (1987:

113), the exine architecture of the non-tectate-

columellate genera of the Pontederiaceae is strikingly

similar (and presumably homologous) to that of mem-

bers of the family Haemodoraceae. The UV-ﬂ uorescent

ferulate in the cell walls, oxalate raphides and styloids,

helobial endosperm with a diminutive chalazal cham-

ber and plasmodial tapetum suggest relationships with

both the Philydraceae and Haemodoraceae (Cook

1998). In addition the seedlings of the deriaceae is very

similar to those of Philydraceae (Tillich 1995).

5. HAEMODORACEAE

R. Brown 1810 (including Conostylidaceae Takhtajan

1987, Dilatridaceae M. Roemer 1840, Wachendorﬁ aceae

W. Herbert 1837, Xiphidiaceae Dumortier 1829).

14/10. Southern Africa, New Guinea, Australia,

Tasmania, eastern and southeastern North America

and Nova Scotia, and tropical America.

5.1 HAEMODOROIDEAE

Mostly with red coloration in roots and rootstocks.

Distinctive tannin cells absent. Starch grains in

vegetative parts simple. Outer cycle of androecium

lacking or (Schiekia) represented by two vestiges of

outer cycle, inner cycle represented by three fertile

stamens or (Pyrrorhiza) by one fertile stamen and

two staminodia. Pollen grains boat-shaped, colpate,

heteropolar, with verrucate or (Schiekia) foveolate

exine sculpturing. Ovary superior or inferior. n = 7,

8, 12, 15, 19–21. – Haemodorum, Lachnanthes,

Dilatris, Schiekia, Pyrrorhiza, Xiphidium, Barberetta,

Wachendorﬁ a.

5.2 CONOSTYLIDOIDEAE

Roots and rootstocks without red coloration. Distinctive

tanniniferous idioblasts present in various organs,

especially in placental tissues. Starch grains in vegeta-

tive parts compound, but those in endosperm simple. In

Superorder COMMELIANAE 717

Phlebocarya silica bodies are present. Stamens six;

staminodia absent Pollen grains, with distinctive rugu-

late (brainlike) exine sculpturing and 2–8 circular

apertures, either fusiform with two apertures

(Anigozanthos, Blancoa, Conostylis spp., Macropidia),

cylindric with two apertures (Phlebocarya), triangular

with three apertures (Conostylis spp.), or globose with

7–8 apertures. Ovary mostly inferior (semi-inferior in

Blancoa). n = 4–8, 11, 14, 16, 21, 28 (mostly 6 or 8). –

Phlebocarya, Tribonanthes, Conostylis, Blancoa,

Anigozanthos, Macropidia.

Related to the Pontederiaceae (Dahlgren and

Rasmussen 1983), which is supported by palynologi-

cal data (Simpson 1987, 1990, 1998), and to the

Philydraceae, supported by such common features as

ensiform leaves, paracytic stomata, the pubescent

ﬂ owering stems, features of helobial endosperm for-

mation and its starchy contents and plasmodial tape-

tum (Hamann 1998), and to both the Pontederiaceae

and Philydraceae in possessing p-coumaric and difer-

ulic acids (Simpson 1998). In addition, members of

Philydraceae and Conostylidoideae and most members

of Pontederiaceae have tannin idioblasts scattered in

various plant organs (Simpson 1998).

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Superorder COMMELIANAE 719

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Order 26. XYRIDALES

Perennial or annual herbs, terrestrial or rarely aquatic

or epiphytic, stemless or caulescent, commonly rhi-

zomatous, rarely bulbous. Vessels in root and stem, or

in all parts (Xyridaceae), with scalariform or more

often with simple perforations. Leaves mostly all at the

base, distichous or spirally arranged, usually with a

sheath, sometimes ligulate, parallel-veined. Stomata

paracytic, anomocytic, seldom tetracytic, hexacytic or

with several subsidiary cells. Flowers solitary or in dif-

ferent type of inﬂ orescences, sometimes much reduced,

bisexual or sometimes (Eriocaulaceae, Hydatellaceae)

unisexual, actinomorphic or zygomorphic, 3-merous,

with or without nectaries. Perianth of six segments

(rarely 4 – Eriopcaulaceae) in two cycles, differenti-

ated into calyx and corolla, rarely (Hydatellaceae)

perianth lacking. Sepals usually free or basally

connate, imbricate or rarely valvate. Petals free or con-

nate, imbricate. Stamens 6 or reduced to 1–3 fertile,

sometimes staminodia present; ﬁ laments short

(Xyridaceae) or slender (Mayacaceae, Hydatellaceae),

adnate to the petals or hypogynous. Anthers mostly

tetrasporangiate and dithecal, or sometimes tetrathe-

cal, or seldom disporangiate and dithecal, basiﬁ xed or

(Eriocaulaceae) dorsiﬁ xed, opening longitudinally

720 Subclass IV. COMMELINIDAE

or by lateral slits (Hydatellaceae) or apical pores.

Tapetum secretory. Microsporogenesis successive or

(Rapateaceae) simultaneous. Pollen grains 2-celled

or seldom 3-celled, 1-colpate or seldom inaperturate,

3-aperturate, spiraperturate, 2-colpate, zonocolpate,

trichotomocolpate, with a pair of short colpae in addi-

tion to the principal one or irregularly colpate,

Gynoecium of three united carpels, seldom one or two

carpels more or less reduced or even suppressed, with

simple or lobed stigma. Placentation central or pari-

etal. Ovary superior, 1–3-locular, sometimes stipitate

(Eriocaulaceae, Hydatellaceae). Ovules solitary to

numerous in each locule, anatropous to hemitropous,

orthotropous or campylotropous, bitegmic, crassinu-

cellate or tenuicellate. Female gametophyte of

Polygonum- or Allium-type. Endosperm nuclear or

helobial, rarely (Hydatellaceae) cellular. Fruits dehis-

cent (loculicidal or rarely septicidal capsules), some-

times indehiscent. Seeds small, sometimes with

embryotega (Mayacaceae); endosperm usually copi-

ous, containing starch and often also protein, and

seldom – oil; embryo small, mostly indifferentiated.

Flavonoids and anthraquinones, ferulic and diferulic

acids present.

Related to the Commelinales and share a common

origin from the commelinalean ancestor.

Key to Families

1 Stamens 6 or 2–3.

2 Flowers solitary or in different types of simple

inﬂ orescences, but not in globose or cylindrical

heads.

3 Leaves mostly all at the base.

4 Leaf base symmetric, silica bodies lacking.

Perennial or less often annual herbs, mostly

of wet habitats (usually marshes, rarely

aquatic), stemless or caulescent. Rootstocks

commonly rhizomes, rarely bulbous. Roots

ﬁ brous, diffuse, mostly ﬁ liform, less often

ﬂ eshy. Raphides are lacking, but small crys-

tals of calcium oxalate present. Vessels in

all parts, with simple perforations often in

oblique or very oblique end walls. Leaves

mostly basal, distichous or less often spi-

rally arranged, ligulate or eligulate, with

short, open sheath, narrow, ﬂ at to terete,

parallel-veined. Stomata paracytic or ano-

mocytic. Inﬂ orescence usually an elongate

or head-like spike, borne at the summit of a

usually nude scape or rarely leafy stem.

Individual ﬂ owers small, bisexual, actino-

morphic to zygomorphic, 3- merous, with-

out nectaries, in the axils of stiff or

coriaceous, spirally arranged, closely imbri-

cate bracts. Perianth of six segments in dis-

tinct two cycles, differentiated into calyx

and corolla; sepals usually free, imbricate,

hyaline or chaffy-scarious, subequal or

more often the outer (anterior) large, more

or less enclosing the rest of ﬂ ower, but

pushed away by the development of corolla,

sometimes more or less reduced or even

obsolete, the lateral ones small, boat-

shaped, keeled; corolla colored, usually

ephemeral, petals free or connate, some-

times to form a rather long, slender tube,

imbricate. Fertile stamens only three and

opposite petals in all genera (with the

exception of one Xyris, which has 6), three

stamens of the outer cycle staminodial or

very rarely absent; ﬁ laments short, adnate

to the petals or hypogynous; anthers tetra-

sporangiate, basiﬁ xed, often elongate,

latrorse or introrse, opening longitudinally.

Microsporogenesis successive. Pollen

grains 2-celled or 3-celled, 1-colpate, 2-col-

pate, irregularly colpate or (Achlyphila,

Abolboda, Aratitiyopea, and Orectanthe)

inaperturate and intectate. Gynoecium of 3

united carpels; style slender, with simple or

3-lobed stigma, sometimes with 2–3 basal

glandular appendages; ovary 3-locular to

1-locular, with several or numerous or

rarely only one ovule per locule or per pari-

etal placenta. Ovules anatropous, orthotro-

pous or weakly campylotropous, weakly

crassinucellate, without parietal cell.

Female gametophyte of Allium- or

Polygonum-type. Endosperm nuclear or

(Abolboda) helobial. Fruits small loculi-

cidal capsules enclosed by the persistent

perianth. Seeds small, numerous, with

strong longitudinal ridges and inner cross-

lines; seed coat formed by both integu-

ments, but only the tegmic layers, which are

ﬁ lled with a resinlike tanninlike substance,

provide mechanical strength; endosperm

copious, rich in aleurone, compound starch

Superorder COMMELIANAE 721

grains, and sometimes also fatty oils;

embryo small, more or less lens-shaped,

little differentiated, lying in the micropylar

part. Anthraquinoses are present. n = 8?, 9,

13, 14, 16, 17 (Xyris).. . . . 2. xyridaceae.

4 Leaf base asymmetric, epidermal cells con-

taining solitary or aggregated silica bodies.

Perennial herbs often of wet habitats, rarely

epiphytic, with short, thick, vertical or pros-

trate rhizome and short stem. Trichomes

nonglandular uniseriate slime-producing

hairs, mucilage cells present. Raphides and

calcium oxalate crystals lacking. Vessels

usually in root and stem, with scalariform

(Saxofriedericioideae) or scalariform and

simple perforations (Rapateoideae). Leaves

crowded in basal rosette, unifacial or bifa-

cial, distichous or less often spirodistichu-

osly arranged, rarely petiolate, with folded

open sheath at base, linear to lanceolate,

often rather ﬁ rm, parallel-veined; ligules

rarely present. Stomata paracytic or less

often tetracytic. Inﬂ orescence axillary or ter-

minal, on leaﬂ ess scapes, mostly with an

involucre of one, two, or several large, coria-

ceous, basally broad bracts subtending the

head or one-sided raceme of spikelets; each

spikelet with a single terminal ﬂ ower and

several imbricate, spirally arranged sterile

bracts beneath it; individual ﬂ owers 3-mer-

ous, bisexual, actinomorphic or nearly so,

with or without nectaries. Perianth segments

six in two cycles strongly differentiated into

calyx and corolla; sepals ﬁ rm, hyaline at

base, free or basally connate, imbricate; pet-

als ephemeral, usually ba-sally connate into

a short or long tube with imbricate lobes.

Stamens six in two cycles; ﬁ laments com-

monly short, usually attached to the corolla

tube and often basally connate; anthers tet-

rasporangiate, basiﬁ xed, linear, usually very

long, sometimes with short, apical connec-

tive appendage, dehiscing by four, two, or

one apical or subapically lateral pores,

extrorse or latrorse. Microsporogenesis

simultaneous. Pollen grains 2-celled, oblong

to elliptic, tectate-columellate, with a thick

foot-layer and granular endexine, 1-colpate

or rarely 2–3-colpate or zonocolpate.

Gynoecium of three united carpels; style

with punctiform or small, capitate and more

or less papillate stigma; ovary 3-locular or

rarely incompletely 3-locular, with one to

several ovules per locule. Ovules anatropous,

crassinucellate, sometimes with conspicuous

funicular obturator, with parietal cell. Female

gametophyte of Polygonum-type. Endosperm

nuclear. Fruits septicidal capsules often with

only one or two fertile locules. Seeds of dif-

ferent shape and sculpturing, with large hilar

scar and a chalazal appendage (caruncle);

seed coat endotestal- exotegmic, formed by

the outer layer of the inner integument and

the innermost of the outer one; endosperm

copious, mealy, rich in simple starch grains

and with a proteinaceous outer layer;

embryo small, lenticular, undifferentiated,

situated at the micropylar end, with ﬂ at side

appressed to endosperm. Accumulate a

considerable amount of aluminium. n = 11

(Maschalocephalus), 26. . .3. rapateaceae.

3 Leaves well distributed along the stem. Small

perennial herbs of shallow fresh water or very

wet habitats with adventitious roots. Raphides,

calcium oxalate crystals, and silica lacking.

Longitudinal septate air canals present in all

vegetative organs. Vessels in root, stem and

leaves, with long scalariform, indistinctly per-

forate end walls. Leaves spirally arranged,

multiranked, not sheathing, commonly with

bidentate apex, 1-veined. Stomata paracytic.

Flowers solitary in the axils of sheathing

bracts, bisexual, actinomorphic, 3-merous,

without nectaries. Perianth segments six, free,

in two distinct cycles; sepals green, valvate to

subvalvate; petals white, imbricate, shortly

clawed. Stamens three, represent the outer

staminal cycle; ﬁ laments slender, glabrous;

anthers tetrasporangiate, basi-ﬁ xed, opening

by apical pores or porelike slits, sometimes by

pores at the end of tubular apical appendage.

Microsporogenesis successive. Pollen grains

2-celled, 1-colpate, columellate, ﬁ nely reticu-

late. Gynoecium of three united carpels; style

ﬁ liform, with capitate or shortly triﬁ d stigma;

ovary 1-locular, with several ovules on each

parietal placenta. Ovules orthotropous, tenuinu-

cellate; micropyle formed by both integuments,

722 Subclass IV. COMMELINIDAE

the inner one closing over the nucellus to form

a narrow endostome. Female gametophyte of

Polygonum-type. Endosperm nuclear. Fruits

loculicidal capsules. Seeds small, ovoid to

globose, minutely beaked and scrobiculate;

seed coat exotestal, mainly formed by the

outer layer of the outer integument, the inner

layer remaining undifferentiated; the inner

layer of the inner integument is tanniniferous,

the outer layer is squeezed ﬂ at (Venturelli and

Bouman 1986); the exotestal cells greatly

increase in size, especially in radial direction

and accumulate starch grains, the lateral and

tangential walls become strongly thickened,

are pitted and contain lignin; these exotestal

cells can hold air and thus instrumental in

keeping the mature seed aﬂ oat for some time

(Venturelli and Bouman 1986); the integu-

mentary layers in the micropylar region are

transformed into a characteristic micropylar

beak, which by many authors (including

myself) are erroneously interpreted as an oper-

culum; it is formed by the inner layer of the

outer integument, the outer layer of the inner

integument and to a lesser extent also by the

inner layer of the inner integument, all of which

undergo an appreciable radial elongation to

close the exo- and endostome; only the cells of

the inner layer of the outer integument have

slightly thickened walls (Venturelli and Bouman

1986); the micropylar tissue probably disinte-

grates before germination and provides a chan-

nel for the emergence of the seedling (Hamann

1961; Venturelli and Bouman 1986); endosperm

copious, the cells of central endosperm are

mainly ﬁ lled with closely-packed simple starch

grains, the outermost layer of the endosperm is

present as an aleurone layer which stores spher-

ical protein bodies and lipids; embryo small,

depressed-ovoid, undifferentiated, apical, lying

opposite micropyle, with ﬂ at side facing the

endosperm, n = 8. . . . . . . . . . 1. mayacaceae.

2 Inﬂ orescence globose to cylindrical, dense, cen-

tripetally ﬂ owering small heads (headlike spikes)

borne on thin peduncles generally supplied with

closed sheath at the base formed by a single leaf,

or composed of many (up to more than 1,000)

such heads: heads subtended by an involucre con-

sisting of more or less numerous, imbricate, vein-

less bracts different in size and shape from bracts

subtending ﬂ owers, although they connected by

intermediates. Small (sometimes very small) to

medium-sized perennial or rarely annual herbs of

various types of habitats – poor sandy soils,

marshy or boggy habitats, rivers, ponds, or tempo-

rarily ﬂ ooded places, sometime even on rocks,

adapted both to short dry periods and to submerged

growth. Perennials with erect or horizontal, often

thick rhizomes. Roots ﬁ brous. Trichomes multi-

cellular, uniseriate, with smooth surface or

glandular, terminal cell simple or bibranched

(malpighiaceous). Raphides lacking, but calcium

oxalate crystals and rarely also silica bodies pres-

ent. Vessels in all vegetative organs, with simple

perforations in roots and simple and scalariform

perforations in stems and leaves. Leaves usually

basal, spirally arranged or rarely distichous, com-

monly linear, parallel-veined, more or less grass-

like, distinctly or indistinctly sheathed at base.

Stomata paracytic. Individual ﬂ owers small or

very small, numerous, sessile or shortly pedicil-

late, unisexual (monoecious or rarely dioecious)

or very rarely bisexual, actinomorphic or slightly

zygomorphic, 3-merous or less often 2-merous,

without nectaries (except for the nectariferous

glands within the tip of the petal in Eriocaulon

and Mesanthemum). Perianth of six or four seg-

ments in two cycles, scarious, often white, dif-

fering in shape and texture but not clearly

differentiated into calyx and corolla; outer seg-

ments membranous- hyaline, free or less often

more or less connate, sometimes with basal tube

or forming spathe; in male ﬂ owers inner segments

usually form 3- or 2-lobed tube, in female ﬂ owers

they are free, often very small or reduced to hairs.

Stamens usually three or two, opposite the petals,

or (Eriocaulon and Mesanthemum) six or four in

two cycles, rarely (Syngonanthus amazonicus)

solitary; ﬁ laments free or rarely connate; anthers

tetrasporangiate or rarely disporangiate, basiﬁ xed

or dorsiﬁ xed, introrse, opening longitudinally.

Microsporogenesis successive. Pollen grains usu-

ally 3-celled, spheroidal to ellipsoid, spirapertu-

rate, echinate to echinulate and may have spinulae

of different size. Male ﬂ owers often with pistillo-

dium. Gynoecium of three or two united carpels;

style often appendaged, 2–3-lobed, the lobes often

biﬁ d at the apex; ovary often stipitate, 3- or 2-loc-

Superorder COMMELIANAE 723

ular, with one ventral-apical, pendulous ovule per

locule. Ovules orthotropous, tenuinucellate, with-

out parietal cell. Female ﬂ owers with conspicuous

staminodia (Rondo-nanthus) or with inconspicu-

ous staminodia or without staminodia. Female

gametophyte of Polygonum-type. Endosperm

nuclear. Fruits small, thin-walled loculicidal cap-

sules. Seeds small, ellipsoidal to subglobose; seed

coat formed by both integuments, but the protect-

ing layer formed primarily by the inner epidermis

of the inner integument; endosperm copious,

mealy, rich in compound starch grains; embryo

very small, lens-shaped or bell-shaped, undiffer-

entiated, situated at the micropylar end. Cell wall-

bound ferulic, p-coumaric, and/or diferulic acids

present. n = 8, 9, 15, 20, 25.. .4. eriocaulaceae.

1 Stamen solitary. Very small aquatic or subaquatic,

annual herbs with numerous unbranched roots and

reduced vascular system. Stem very short, branch-

ing to form a leafy tuft, often with minute multicel-

lular hairs in leaf axils. Oxalate raphides and silica

bodies lacking. Vessels in roots and stem, with sca-

lariform perforations. Leaves all basal, submerged

and emergent, alternate, glabrous, linear to ﬁ liform,

lack a distinct sheath or ligules, 1-veined. Stomata

anomocytic (Trithuria), in submerged species of

Hydadella apparently lack stomata (Hamann 1998).

Inﬂ orescence terminal, sessile or scapose head with

an involucrum of 1–4 pairs of opposite, membra-

nous, 1-veined bracts often forming an apparent

whorl; ﬂ owers numerous, minute, unisexual, lack-

ing perianth and bracteoles, hydrophilous or autog-

amous. Filaments ﬁ liform; anthers tetrasporangiate,

basiﬁ xed, linear to elliptic, opening longitudinally

by lateral slits. Pollen grains 2-celled, tectate-colu-

mellate, 1-colpate and spinulose (Trithuria) or with

indistinct porelike aperture. Gynoecium monomer-

ous or more probably pseudo-monomerous (accord-

ing to Hamann, 1998, the three prominent ribs on

the ovary wall in Trithuria and the 3-valved fruit in

T. submersa indicate a basically 3-merous gynoe-

cium); stigma terminal, sessile, with 2–10 hairs,

each a single row of distended cells; ovary superior,

shortly stipitate, 1-locular, with one pendulous

ovule. Ovules anatropous, crassinucellate or

pseudocrassinucellate. Endosperm very probably

cellular (Hamann 1998). Fruits small, non-ﬂ eshy,

dehiscent or indehiscent, 3-valvular capsules

(Trithuria), or achenelike (Hydatlla). Seeds minute,

ovoid, with an operculum formed by the apical cells

of the inner layer of the inner integument; seed coat

includes both testa and tegmen, but the protective

layer represented by derivatives of the outer epider-

mis of the outer integument (seeds exotestal); most

part of endosperm replaced by starchy perisperm, the

rest of endosperm occupies small zone close to

embryo and consists of a few cells devoid of starch;

embryo minute, peripheral, lens-shaped, very weakly

differentiated. . . . . . . . . . . . . . . .5. hydatellaceae

1. MAYACACEAE

Kunth 1842. 1/4 or up to 10. Southeastern United

States, West Indies, Central and South America, 1

species in Africa (Angola, Zaire, Zambia).

Mayaca.

They share more characters with the Xyridaceae.

According to Stevenson (1983, 1998), Venturelli and

Bouman (1986) and Johri et al. (1992), embryologi-

cally Mayacaceae resemble the Xyridaceae more than

the Commelinaceae and can be placed near Xyridaceae.

However they markedly differ from the Xyridaceae in

aquatic habit, anatomy of vegetative organs, dispersed,

1-veined leaves without distinct sheath, solitary ﬂ owers,

apically dehiscing anthers, pollen morphology.

2. XYRIDACEAE

C. Agardh 1823 (including Abolbodaceae Nakai

1943). 5/325. Widespread in tropical and subtropical

regions, especially of America, with relatively few

species extending into temperate regions. All ﬁ ve gen-

era have representatives on the Guiana Highlands,

which might be the primary centre (Kral 1998).

Very close to the Mayacaceae and share a common

origin from the commelinalean ancestor.

2.1 XYRIDOIDEAE

Leaves more or less clearly distichous, unifacial, often

with an abrupt transition fro sheath to lamina. Petals

commonly free, yellow. Pollen grains smooth and

1-colpate (occasionally 2-colpate). Staminodia typi-

cally present and biﬁ d (lacking in Achlyphila). Styles

symmetric and unappendaged. Ovary 1-locular or

incompletely 3-locular, with parietal or free basal

placenta. – Achlyphila, Xyris.

724 Subclass IV. COMMELINIDAE

2.2 ABOLBODOIDEAE

Leaves polystichous, bifacial, without abrupt transi-

tion from sheath to lamina. Petals united, blue or white.

Pollen grains spinulose and inaperturate. Staminodia

reduced to ﬁ laments or lacking. Styles mostly asym-

metric and appendaged. Ovary 3-locular, with axile

placentas. – Abolboda, Orectanthe, Aratitiyopea.

3. RAPATEACEAE

Dumortier 1829. 16/80. Mainly tropical South

America, especially Guyana Highlands, with one

genus (Epidryos) extending into Panama and one

monotypic genus (Maschalocephalus) in tropical West

Africa (Sierra Leone and Liberia).

3.1 SAXOFRIDERICIOIDEAE

Carpels with several ovules, on axil or on septa. Seeds

prismatic or pyramidal. – saxofridericeae:

Saxofridericia, Phelpsiella, Stegolepis, Amphiphyllum,

Marahuacaea, Epidryos; sphoenocephaleae:

Kunhardtia, Guacamaya, Schoenocephalium.

3.2 RAPATEOIDEAE

Carpels with one ovule inserted at or near the base.

Supernumerary antipodals are common. Seeds oval or

oblong, with ﬂ attened and papillate apical appendages. –

RAPATEAE: Rapatea, Cephalostemon (including

Duckea), Spathanthus; monotremeae: Monotrema,

Potarophytum, Windsorina, Maschalocephalus.

A very heterobathmic family with relatively primi-

tive vessels in many genera (Carlquist 1966, 1969)

and a rather specialized inﬂ orescence and ﬂ owers.

The Rapateaceae are usually placed near the

Xyridaceae (see Hallier 1912; Hutchinson 1934,

1959, 1973; Engler’s Syllabus 1964; Takhtajan 1966,

1980, 1987; Cronquist 1980, 1988; Dahlgren et al.

1985; Thorne 1992a, b). However, there are many

important differences, including asymmetric leaf

bases, the presence of silica bodies in the epidermal

cells and tannin- bearing cells, the peculiar chloren-

chyma cells, and the nature of the vascular core of the

roots (Tomlinson 1969), poricidal dehiscence of

anthers, pollen morphology, simultaneous microsporo-

genesis, as well as morphology of inﬂ orescences,

fruits, and seed ana-tomy, and accumulation of alumi-

num. Tiemann (1985) is of the opinion that the

Rapateaceae should be placed closer to the Cyperaceae

and Juncaceae to which they are related because of

numerous common embryological features. Also in

Tiemann’s opinion ﬂ ower morphology and anatomy

support a relationship with the Thurniaceae, which

she considers a connecting link. However, the

Rapateaceae differ markedly from the Juncales in

their pollen grains and seed coat structure as well as

in their perianth morphology and some anatomical

features. Many years ago Pilger (1930) and Smith

(1934) suggested the relationship of the Rapateaceae

with the Bromeliaceae. However, morphologically

the Rapateaceae are nearer to the families of the

Commelinanae, especially to the Xyridaceae.

4. ERIOCAULACEAE

Martynov 1820. 10/1000-1500. Mainly tropical

regions, especially of America; a few species of

Eriocaulon extend into temperate regions of Europe,

eastern Asia and North America and most species

Lachnocaulon occur in United States.

4.1 ERIOCAULOIDEAE

Submerged and ﬂ oating, rarely terrestrial plants. Roots

and leaves with aerenchyma. Petals free, with black

tips; stamens adnate to the corolla. – Eriocaulon.

4.2 PAEPALANTHOIDEAE

Usually terrestrial plants. Petals often connate. Stylar

appendages, stigmas commissural. – Rondonanthus,

Mesan themum, Lachnocaulon, Syngonanthus, Philodice,

Leiothrix, Tonina, Paepalanthus, Blastocaulon.

A distinct and rather advanced family that has some

similarities with the Xyridaceae, including many veg-

etative characters, compact inﬂ orescences, echinate

pollen, and stylar appendages like those in some

Xyridaceae. However, the Eriocaulaceae differ from

the Xyridaceae in unique type of inﬂ orescences, spi-

raperturate pollen grains, solitary orthotropous pendu-

lous ovule per locule, endotegmic seeds.

5. HYDATELLACEAE

Hamann 1976. 2/10. Temperate Australia and

Tasmania, 1 species of Hydatella in New Zealand

and 1 species of Trithuria in India.

Hydatella, Trithuria.

Superorder COMMELIANAE 725

A very specialized and isolated family which related

to Xyridaceae and Mayacaceae (Bremer 2002;

Soltis 2006).

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