Takhtajan Armen. Flowering Plants

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706 Subclass IV. COMMELINIDAE

irregularly, occasionally indehiscent (nuts or

achene-like). Seeds arillate, ﬁ lled with strongly

developed perisperm containing copious starch

composed of simple concentric starch grains and

well-developed chalazal tissue; endosperm is

little developed and contains starch; embryo

cylindrical, rather short, has a cotyledonary

sheath and contains fat; testa encrusted with

phytomelan. Benzoquinones and steroidal

saponins present. n = 8, 9, 14. . . .6. costaceae.

5 Pollen grains spinulose. Perennial rhizoma-

tous herbs, sometimes up to 5 m tall. Stem and

rhizome with conspicuous mucilage canals or

cavities. Vessels only in root, with simple per-

forations, or roots without vessels. Starch

grains ﬂ attened. Leaves distichous to spirally

arranged, with arching-convergent lateral

veins and without pulvinus; the mesophyll

containing calcium oxalate crystals. Stomata

paracytic, tetracytic or hexacytic. Flower

bisexual, in terminal spikes or in panicles

(thyrses), usually composed of 2-ﬂ owered cin-

cinni, pairs not enantiomorphic. Inner stamin-

odia petaloid. The single fertile stamen bearing

only one half-anther; anther introrse, dispo-

rangiate, unilocular. Style ﬂ at and ﬂ eshy-peta-

loid, with terminal stigma. Ovary 3-locular;

ovules 20–50 per locule, anatropous. Fruits

warty capsules, opening by collapse of the

pericarp. Seeds exotestal, exarillate, with

starch, without operculum but with distinct

germination slit and micropylar collar;

perisperm without canals; embryo straight;

endosperm consisting of a thin, starchy layer.

Present cyanidin, ﬂ avonols (kaempferol and

quercetin), chelidonic acid and aromatic resin;

n = 9.. . . . . . . . . . . . . . . . . . . .7. cannaceae.

5 Pollen grains psilate. Few-ﬂ owered herbs,

rosulate or with aerial stem of variable length,

sometimes lianescent and highly-climbing,

rarely epiphytic. Mucilage canals usually

lacking (but present in Thalia). Vessels in

stems and roots with simple or scalariform

perforations. Starch grains ellipsoidal and

slightly eccentric, rarely ﬂ attened or angular.

Leaves generally distichous, with arching

lateral veins that fuse marginally, some of

them at intervals being thicker than the oth-

ers; petiole with swollen pulvinus at or near

the summit that controls the orientation of

lamina. Flower pairs are in mirror image

(enantiomorphic). The single fertile stamen

with a pollen sac on one edge). Two inner

staminodia (cucullate and callous-thickened)

modiﬁ ed into structures for the explosive

release of pollen. Septal nectaries present.

Style not petaloid, often expanded and lobed

at the summit, with lateral stigma in depres-

sion between lobes. Ovary 3-locular, but two

often empty or absolete; ovules solitary in

fertile locule, anatropous or anacampylotro-

pous Fruits commonly 1- to 3-seeded cap-

sules with dry to ﬂ eshy pericarp, sometimes

baccate or (Thalia) caryopsis-like. Seeds

rather large, endotestal, usually with a basal

aril (lacking or transformed into a pulp in

indehiscent fruits) and with an operculum;

testa encrusted with phytomelan; embryo

horseshoe-shaped; perisperm with simple,

laterally dilated or variously branched canals

that originates during seed maturation by an

intrusion of the chalazal tissue into the

nucellus (Grootjen 1983). Proanthocyanidins

(cyanidin) and ﬂ avonols (kaempferol and

quercetin) present or absent; n = 4–14, 21,

23. . . . . . . . . . . . . . . . . . . 8. marantaceae.

1. STRELITZIACEAE

Hutchinson 1934. 3/7. Tropical South America (Phe-

nakospermum), East coast of South Africa from Cape

Province to Transvaal (Strelitzia), and Madagascar

(Ravenala).

ravenaleae: Ravenala; strelitzieae: Phena-

kospermum, Strelitzia.

According to Tomlinson (1962, 1969), the Strelit-

ziaceae may be considered to represent the nearest

approach to the ancestral stock from which the whole

order Zingiberales s.1. has evolved. However,

Strelitziaceae are a very heterobathmic group that

combines some primitive characters (such as complete

or nearly complete androecium) with some very

advanced characters (such as very specialized perianth

and inferior ovary). I agree with Kress (1990: 716) that

“the arborescent nature of the Strelitziaceae is a derived

character and not a primitive feature as (Tomlinson

1962) suggested.”

Superorder ZINGIBERANAE 707

2. LOWIACEAE

Ridley 1924. 1/16. Southern China (Kwangtung prov-

ince and Hainan), Indochina, Malay Peninsula,

Sumatra, Java, Borneo, and Brunei.

Orchidantha (Lowia).

A very distinct taxon characterized by many unique

characters that isolate it from other zingiberids (see Kress

1990). However, it deﬁ nitely belong to the “banana

group” of the Zingiberanae (Kress 1995) and has many

common features with the Musaceae (including opercu-

late seeds) and especially with the Strelitziaceae ( including

capsular fruits, arillate seeds and pollen grains resembling

much of Ravenala) (Larsen 1998).

3. MUSACEAE

A. L. de Jussieu 1789. 3/60. Tropical Africa (from the

Cameroon to Ethiopia and Transvaal), India, south-

eastern Asia, Malesia, New Guinea, northeastern

Australia, and the Paciﬁ c Islands.

Musa, Ensete, Musella.

4. HELICONIACEAE

Nakai 1941. 1/200 +. Southern Central America and

South America (especially northern parts of the Andes),

with a few species in, Melanesia.

Heliconia.

Markedly differ from the related Musaceae in the

absence or articulated laticifers, root anatomy, distic-

hous leaves, bisexual ﬂ owers, adaxially placed median

member of the outer cycle of the perianth, the presence

of a single staminodium opposite the unpaired adaxial

member of the outer cycle of perianth, plasmodial

tapetum, and especially the unique type of the schizo-

carpous fruits.

5. ZINGIBERACEAE

Martynov 1820 (including Alpiniaceae Link 1821).

53/1350. Madagascar, Mascarenes, Seychelles, tropical

and South Africa, southern, eastern, and Southeast

Asia, Malesia, Paciﬁ c Islands, New Guinea, and north-

ern and eastern Australia (mainly Queensland), with

some representatives in Neotropics. The center of

diversity is in tropical Asia. Tamijia – Kalimantan,

Siphonochilus – Africa and Madagascar.

Classiﬁ cation after Kress, Prince and Williams

(2002).

5.1 TAMIJIOIDEAE

1/1. Plants evergreen with ﬁ brous rhizomes; ovary uni-

locular with parietal placentation. Plane of distichy of

leaves perpendicular to rhizome. Lateral staminodes

well developed and fused to labellum. – Tamijia.

5.2 SIPHONOCHILOIDEAE

1/15. Plants with seasonal dormancy period and ﬂ eshy

rhizomes; ovary trilocular with parietal placentation.

Plane of distichy of leaves perpendicular to rhizome.

Lateral staminodes well developed and fused to

labellum. – Siphonochilus.

5.3 ALPINIOIDEAE

Lateral staminodes reduced or absent. Plane of distichy

of leaves perpendicular to rhizome. – alpinieae: Afra-

momum, Alpinia, Amomum, Autolandra, Cyphos-tig ma,

Elettaria, Elettariopsis, Etlingera, Geocharis,

Geostachys, Hornstedtia, Leptosolena, Paramomum,

Plagiostachys, Renealmia, Vanover berghia; riedelieae:

Burbidgea, Pleurantho dium, Riedelia, Siamanthus.

Insertae sedis: Siliquamomum (1, southern China,

Indochina).

5.4 ZINGIBEROIDEAE

Plane of distichy of leaves parallel to rhizome. –

zingibereae: Boesenbergia, Camptandra, Cautleya,

Cornukaempferia, Curcuma, Curcumorpha, Dis-

tichochlamys, Hanifﬁ a, Haplochorema, Hedychium,

Hitchenia, Kaempferia, Laosanthus, Nanochilus,

Para cautleya, Parakaempferia, Pommereschea, Pyrgo-

phyllum, Rhynchanthus, Roscoea, Sca phochlamys,

Smithatris, Stadiochilus, Stahlianthus, Zingiber; glob-

beae: Gagnepainia, Globba, Hemiorchis, Mantisia.

Insertae sedis: Caulokaempferia (10, Himalayas,

Southeast Asia).

6. COSTACEAE

Nakai 1941. 4/120. The largest genus Costus (95) pan-

tropical, Monocostus (1, Peru) and Dimerocostus (3–5)

neotropical, Tapeinochilus (18) in East Malesia to

northeastern Australia.

708 Subclass IV. COMMELINIDAE

Costus, Dimerocostus, Monocostus, Tapeinochilos.

Close related to the Zingiberaceae.

7. CANNACEAE

A. L. de Jussieu 1789. 1/10-25. Tropical and subtropi-

cal regions of America, especially of western South

America.

Canna.

Related to the Marantaceae. “Differences between

the two families including the positioning of pollen on

the ventral face in Cannaceae, the dorsal face in

Marantaceae, indicate that both are different lineages”

(Kubitzki 1998).

8. MARANTACEAE

R. Brown 1814. 31/550. Nearly pantropical, but espe-

cially numerous in tropical America and absent in

Australia.

Phrynium, Ataenidia, Monophrynium, Phacelo-

phrynium, Stachyphrynium, Galathea, Sanblasia, Pleios-

tachya, Ischnosiphon, Monotagma, Donax, Schumanni-

anthus, Hypselodelphys, Trachyphrynium, Megaphrynium,

Sarcophrynium, Maranta, Maran tochloa, Afrocalathea,

Monophyllanthe, Koernickanthe, Myrosma, Saranthe,

Hylaeanthe, Ctenanthe, Stromanthe, Halopegia, Thalia,

Thaumato coccus, Cominsia, Haumania.

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University, China.

Superorder COMMELIANAE

Order 25. COMMELINALES

Perennial or rarely annual, mostly terrestrial herbs with

various types of underground storage organs, including

rhizomes, root tubers or, rarely, bulbs. Raphides (cal-

cium oxalate crystals) are present in raphide sacs or

raphide canals; silica bodies present in some genera of

Commelinaceae and Hanguana. Vessels in roots, some-

times also in stems and leaves, with scalariform of sim-

ple perforations. Leaves alternate, spirally arranged or

distichous, petiolate or sessile, basal or cauline, simple,

entire, usually with parallel venation. Stomata usually

tetracytic or paracytic. Flowers in various kinds or termi-

nal or axillary inﬂ orescences, sometimes solitary, bisex-

ual or rarely unisexual, actinomorphic to zygomophic,

usually 3-merous, bracteate or not. Perianth segments

six in two cycles, rarely (Philydraceae) of four perianth

segments. Stamens basically six in two cycles, rarely

stamen solitary (Philydraceae), or three stamens, with or

without staminodia; ﬁ laments free or basally connate,

glabrous or sometimes pubescent; anthers basiﬁ xed or

dorsiﬁ xed, introrse to extrorse, sometimes versatile, tet-

rasporangiate, usually opening longitudinally. Tapetum

secretory, amoeboid. Microsporogenesis successive.

Pollen grains generally 2-celled and 1- colpate, some-

times inaperturate. Gynoecium of three united carpels;

style with capitate, 3-lobed stigma; ovary mostly 3-locu-

lar, superior or rarely inferior, with solitary to numerous

ovules per locule. Ovules anatropous, hemitropous,

orthotropous, or rarely campylotropous, bitegmic,

crassinucellate to tenuinucellate. Female gametophyte

of Polygonum-type or rarely Allium-type. Fruits mostly

loculicidal capsules, rarely indehiscent and than some-

times baccate. Seeds usually small, of various shapes

and sculpturing, rarely arillate; embryo small, usually

straight, well- differentiated to undifferentiated;

endosperm copious, with starch grains and aleuron bod-

ies. Containing ﬂ avonoids, ferulic and diferulic acids.

Have many similarities with the Bromelianae as

well as with the Zingiberanae. They are linked with the

Bromelianae and Zingiberanae by such conspicuous

synapomorphies as epiticular wax of the Strelitzia-

type, cell walls containing UV-ﬂ uorescent compounds,

and endosperm with copious starch (Dahlgren and

Rasmussen 1983).

712 Subclass IV. COMMELINIDAE

Key to Families

1 Vessels in all organs. Perennial or sometimes

annual, terrestrial or rarely epiphytic herbs with rhi-

zomes, root tubers or, rarely, bulbs. Roots ﬁ brous or

tuberous. Calcium oxalate crystals are present in

raphide sacs forming articulated raphide-canals

(lacking only in Cartonema). Contain silica bodies.

Starch, when present in vegetative parts, consists of

simple, excentric grains. Vessels usually in all

organs, with simple perforations. Three-celled glan-

dular microhairs present in most genera. Leaves

usually alternate, spirally arranged or distichous,

simple, entire, commonly somewhat succulent,

often narrowed into a false petiole, parallel-veined;

leaf-sheath closed and well- deﬁ ned. Stomata tetra-

cytic, hexacytic, or sometimes paracytic. Flowers in

various kinds of terminal or axillary, basically

cymose inﬂ orescences, sometimes solitary, bisexual

or sometimes polygamous, actinomorphic or more

or less zygomorphic, 3-merous, without nectaries.

Perianth segments six in two distinct cycles. Sepals

green or sometimes petaloid and colored, usually

free, imbricate, generally similar. Petals ephemeral,

free and sometimes clawed or occasionally basally

connate, imbricate. Stamens basically six in two

cycles, but often there are only three to one fertile

stamens and two to four staminodia, rarely three or

fewer fertile stamens without staminodia; ﬁ laments

slender, free or rarely basally connate, glabrous or

some or all bearded with brightly colored long

hairs; anthers basiﬁ xed or dorsiﬁ xed, sometimes

versatile, introrse to extrorse, opening longitudi-

nally or rarely by apical pores or porelike slits.

Tapetum amoeboid. Pollen grains generally

2-celled, mostly 1-colpate, rarely with two addi-

tional apertures. Gynoecium of three united carpels;

style with capitate, 3-lobed or penicillate stigma;

ovary sessile or stipitate, 3-locular or rarely 2-locu-

lar (1 locule being underdeveloped or even sup-

pressed), with 1 to ca. 20 ovules per locule. Ovules

anatropous, hemitropous, orthotropous, or rarely

campylotropous, crassinucellate to tenuinucellate,

with or without parietal cell. Female gametophyte

of Polygonum-type or rarely Allium-type.

Endosperm nuclear. Fruits mostly loculicidal cap-

sules, rarely indehiscent and then sometimes bac-

cate. Seeds generally small, hard, of various shapes

and sculpturing, rarely arillate; seed coat testal-teg-

mic, formed by endotesta and exotegmen and in

some degree also endotegmen (Vyshenskaya 1985),

with disclike or conic operculum (embryotega) cov-

ering the embryo and surrounded by a micropylar

collar except in Cartonema, where it still can be

identiﬁ ed as a bump; endotesta siliciﬁ ed; endosperm

copious, starchy, contains compound starch grains

and a small amount of aleurone; embryo small,

straight, lateral, well differentiated except in

Cartonema, situated at the opposite end to the hilum

beneath the operculum. 6-hydroxyﬂ avonoids are

present. n = 4 to 29.. . . . . . . . .2. commelinaceae.

1 Vessels only in roots and sometimes in stems.

2 Vessels with scalariform perforation

3 Ovules crassinucellate

4 Tapetum secretory, its cells 2–4-nucleate and

always contain raphids. Perennial pubes-

cent herbs with rhizomes or corms and

ﬁ brous roots. Small glandular hairs and

longer, uniseriate hairs often present.

Tanniniferous cells and idioblasts with sty-

loids of calcium oxalate widespread in all

organs, but silica bodies lacking in the veg-

etative parts. Vessels in roots and in stems

(Schnaider and Carlquist 2005), with sca-

lariform perforations. Leaves sessile, unifa-

cial, ensiform to linear or terete, mostly

basal, basal and lower cauline leaves distic-

hous, with well-developed sheath, the rest

smaller and spirally arranged; venation par-

allel. Stomata paracytic, but in Helmholtzia

and Orthothylax have two to four lateral

and two less distinct polar subsidiary cells.

Flowers in terminal simple or paniculately

branched spikes, sessile in the axils of rather

large spathaceous bracts, bisexual, strongly

zygomorphic, 3-merous. Perianth petaloid,

of four free or basally connate members;

the large (adaxial) member, which is fre-

quently 3- or 2-cuspidate, consists of

3 fused perianth segments – 2 segments of

the outer cycle and the reduced median

(adaxial) segment of the inner cycle; two

other lateral segments of the inner cycle are

very small, resemble staminodia, and are

basally more or less adnate to the stamen;

the large abaxial member of the perianth

represents the only free segment of the

Superorder COMMELIANAE 713

outer cycle (Hamann 1966, 1998, 1998).

Stamen solitary, opposite the abaxial peri-

anth segment, corresponds to the median

stamen of the outer cycle of the ancestral

androecium which is otherwise suppressed;

the inner cycle of stamens is always lack-

ing; ﬁ lament strongly ﬂ attened, inserted at

the base of the perianth segment; anthers

basiﬁ xed to dorsiﬁ xed, opening longitudi-

nally, straight and introrse, or curved

(Phillydrella), or helically twisted

(Philydrum). Tapetum secretory, its cells

2–4-nucleate and always contain raphids.

Pollen grains 2-celled, in monads or

(Philydrum) tetragonal tetrads, 1-colpate,

tectate-columellate, with distinctive lamel-

lae inside the homogeneous foot layer,

reticulate; the aperture wall consists of a

thick, 2-layered intine with exine absent or

composed of scattered deposits (Simpson

1985). Gynoecium of 3 united carpels; style

persistent, with a capitate, slightly 3-lobed

stigma; ovary superior, 3-locular or for the

greater part 1-locular with deeply intruded

and bibrachiate parietal placentas, with

numerous ovules on each placenta. Septal

nectaries or other nectariferous glands

absent. Ovules anatropous, weakly crassi-

nucellate, with parietal cell. Female game-

tophyte of Polygonum-type. Endosperm

formation of a peculiar subtype of the helo-

bial type, in which the small chalazal

endosperm chamber becomes cellular ear-

lier than the large micropylar chamber; only

the micropylar chamber serves as storage

tissue (Hamann 1966, 1998, 1998). Fruits

loculicidal capsules or rarely berrylike.

Seeds small to minute, generally very

numerous, reddish-brown, ellipsoidal to

ovoid or linear-cylindrical, spirally striate-

tuberculate (Philydrum), reticulate-rugose

(Philydrella) or silky-striate (Helmholtzia),

with generally conspicuous red-colored

chalazal cap and an operculum (later pushed

off by the germinating embryo) formed by

the micropylar region of both layers of the

inner integument; seed coat testal-tegmic;

endosperm copious, rich in starch (consist-

ing of large single grains), crystalline aleu-

ron bodies and also some oil; embryo small,

linear, straight, weakly differentiated.

Ferulic and diferulic acids present in the

cell wall, also contain proanthocyanins. n =

8, 16, 17.. . . . . . . . . . . . . 3. philydraceae

4 Tapetum amoeboid; cells 1–2-nucleare

without raphids. Perennial or rarely

(Hydrothrix) annual, rhizomatous or stolo-

niferous, swamp or aquatic herbs ranging

from amphibious through ﬂ oating-leaved

and free-ﬂ oating to almost completely

submerged. Branch ing of the main axis

usually sympodial. Trichomes are absent

on vegetative organs. Tanniniferous idio-

blasts have been reported from the peti-

oles. Oxalate raphides are widely present

and cells with one or two prismatic crys-

tals have been found in some genera.

Vessels in roots and often also in stem,

with scalariform perforations. Leaves both

basal and cauline, distichous or spiro-dis-

tichuous along stem, usually with consid-

erable sheath enveloping stem, with

distinct and sometimes inﬂ ated petiole and

lanceolate to ovate or broadly cordate,

rarely linear or even; ﬁ liform undifferenti-

ated lamina, sometimes reduced. Stomata

paracytic. Flowers in terminal racemes,

spikes, or panicles or solitary, often sub-

tended by two spathelike bracts, bisexual,

actinomorphic to zygomorphic, 3-merous,

bracteate or not, sometimes triheterostylous

and with trimorphic pollen grains. Perianth

of 6 (in Scholleropsis mostly 4) usually

petaloid segments in two cycles, nearly

free or more often basally connate into a

tube. Stamens mostly six in two cycles or

only three stamens (sometimes unequal),

rarely (Scholleropsis) three stamens and

one staminodium, or (Hydrothrix) only

one stamen and two staminodia; ﬁ laments

slender, attached to the perianth tube;

anthers basiﬁ xed or peltate, opening longi-

tudinally and introrse or opening by apical

pores. Tapetum amoeboid, cells 1–2-nucle-

ate. Pollen grains 2-celled or rarely

3-celled, with 1–3 distal or subequatorial

714 Subclass IV. COMMELINIDAE

colpi, exine verrucate, scabrate to ﬁ nely

reticulate. Gynoecium usually of three

united carpels (in Pontederia pseudomono-

merous); style slender, with capitate or

3-lobed stigma; ovary superior, mostly

3-locular, with several or numerous ovules

or less often (Pontederia) with one apical

pendulous ovule. Septal nectaries mostly

present. Ovules anatropous, crassinucel-

late, with parietal cell. Female gameto-

phyte of Polygonum-type. Endosperm

helobial, with a small chalazal chamber

where cell wall formation begins much

earlier than in the large micropylar cham-

ber. Fruits many-seeded loculicidal cap-

sules or (Pontederia and Reussia) 1-seeded

nutlets surrounded by postﬂ oral-modiﬁ ed,

hardened, rigid perianth bases. Seeds

small, usually longitudinally furrowed,

with relatively large, cylindrical embryo

surrounded by copious endosperm consist-

ing of elongate cells containing starch and

aleurone layer; cotyledon linear, bifacial;

seed coat exotestal; exotestal cells box-

like, elongated, endotestal cells thin, elon-

gated transversely; the inner one is

represented by strongly compressed layer

containing a reddish-brown pigment. n =

(7)8(-13). . . . . . . . . . 4. pontederiaceae

3 Ovules tenuinucellate. Robust perennial herbs

growing in humid forests, lowland swamps or

along slowly moving rivers and lakes (some-

times ﬂ oating), with creeping rhizomes and

long stolons arising from the base of the erect

stems and forming dense colonies. Short multi-

cellular branched hairs common on stem and

leaf of young plants, each with a deeply

sunken, uniseriate base. Conspicuous tannin

cells present. Vessels only in roots, with scalari-

form perforations. Leaves rolled unilaterally in

bud (ptyxis supervolute), mostly basal, spirally

arranged, linear to lanceolate, acute, pseudo-

petiolate, with strong midrib, parallel longitudi-

nal veins are linked with numerous close-set

thin cross veinlets, sheathing at base, usually

with silica bodies; the sheat and petiolar region

contain large mucilage channels. Stomata tetra-

cytic. Flowers small, in large terminal panicles

composed of spike-like partial inﬂ orescences,

with broad, large bracts, dioecious, actinomor-

phic, 3-merous. Perianth segments six in two

cycles, shortly connate at base, persistent, the

inner cycle red-dotted, larger and vaulted. Male

ﬂ owers with rudimentary gynoecium which is

surrounded by a lobed, disc-like structure

Stamens six, inserted on the base of perianth

segments; ﬁ laments subulate with broader base;

anthers small, basiﬁ xed, tetrasporangiate, open-

ing longitudinally, introrse; female ﬂ owers with

six staminodia. Tapetum amoeboid, with a

periplasmodium. Pollen grains more or less

spheroidal, inaperturate, with thin, hyaline

exine provided with small spinules. Gynoecium

of three united carpels; ovary superior, 3-locu-

lar, with one ovule per locule; stigma subsessile

or sessile, broadly 3-lobed. Characteristic

mucilage-secreting hairs present within the

ovary locule (Rudallet al. 1999). Ovules almost

orthotropous, probably tenuinucellate, with

funicular obturator. Fruits ﬂ eshy thick-walled

1–3-seeded drupes. Seeds bowl-shaped, meso-

endotestal, without phytomelan or phloba-

phene; endosperm copious, clearly divided into

several (up to 7) separate segments which pos-

sibly are the remnants of nucellus (Nemirovich-

Danchenko 1985) and enclosed by a single

peripheral later of aleurone cells, with simple

and compound starch grains of various size and

form (Nemirovich-Danchenko); embryo small,

globose. n = 12, 18, and more than 45; chromo-

somes very small.. . . . . . . . 1. hanguanaceae

2 Vessels with simple or rarely scalariform

perforation. Perennial rhizomatous and stolonif-

erous or less often cormose to bulbous herbs

mostly with various kinds of trichomes on both

the vegetative and reproductive parts. Roots fas-

ciculate and ﬁ brous, sometimes tuberous. Oxalate

raphides present in all parts, but silica bodies

lacking. Sieve-element plastids of form-P2c.

Vessels usually only in roots but sometimes

(Lachnanthes) also in stem, with simple or

(Dilatris and Tribonanthes, Anigozanthos ﬂ avi-

dus – Conostyl idaceae) scalariform perforations.

Leaves mostly basal or subbasal, distichous, lan-

ceolate, narrowly linear or acicular, unifacial,

sheathing at base, parallel-veined, sometimes

(Barberetta and Wachendorﬁ a) plicate. Stomata

paracytic. Flowers in terminal, generally cymose

Superorder COMMELIANAE 715

inﬂ orescences or solitary, pedicellate or subsessile,

bisexual, actinomorphic to strongly zygomorphic,

3-merous, conspicuously hairy or glabrous.

Perianth segments six, persistent, membranous to

coriaceous, either composed of three outer and

three inner imbricate, free or basally connate

similar segments or (in Anigozanthos, Blancoa,

and Conostylis) perianth connate into a short or

long, straight or curved tube with six monocyclic

and valvate lobes. Stamens six in two cycles or

three opposite the inner perianth segments or

(Pyrrorhiza) one opposite the inner anterior peri-

anth segment and two staminodia opposite the

inner latero-posterior segments; ﬁ laments free or

adnate to perianth tube, equal or unequal in length

and/or position; staminodia sometimes present;

anthers generally oblong, tetrasporangiate, bas-

iﬁ xed to dorsiﬁ xed, opening longitudinally,

introrse, straight or arcuate, sometimes with a con-

nective appendage. Tapetum amoeboid; radially

oriented, ligniﬁ ed cell wall thickenings occur in

the endothecial and middle layer cells (Simpson

1998b). Pollen grains 2-celled, with 1–3-layered

atectate exine, 1-colpate or 2–8-porate. Gynoecium

of three united carpels, all fertile or (Barberetta)

only the posterior carpel fertile at maturity; style

slender, straight to strongly curved to one side of

the ﬂ ower, with capitate or 3-lobed stigma; ovary

superior to inferior, 3-locular or (Phlebocarya)

basally 3- and apically 1-locular, with one to

numerous ovules per carpel. Septal nectaries pres-

ent or rarely (as in Xiphidium) absent. Ovules ana-

tropous or orthotropous, crassinucellate, with

parietal cell. Female gametophyte of Polygonum-

type. Endosperm helobial, with early cell forma-

tion in the chalazal chamber. Fruits loculicidal to

apically poricidal capsule, sometimes indehiscent

(in Macropidia dehiscing into three single-seeded

mericarps with a hard endocarp). Seeds of various

shapes, glabrous or hairy; seed coat formed by

both integuments, either predominantly by the

outer integument (in Haemodoroideae) or mainly

by the inner integument (Conostylidoideae);

embryo globose to ovoid or kidney-shaped, lateral,

in Conostylidoideae minute and undifferentiated,

in Haemodoroideae larger and more differenti-

ated; endosperm copious, differentiated into

peripheral aleurone and lipid layer, in which cell

walls store cellulose, and an inner layer containing

simple or compound starch grains. Flavones, fruc-

tans and arylphenalenones are present. The order

is chemically unique in possessing perinaphthe-

none pigments which are responsible for the red

coloration prominent in the roots and rootstocks of

the Haemodoraceae s. str. Chelidonic acid widely

distributed in the order. Species of Wachendorﬁ a

and Anigozanthos have cell wall-bound p-cou-

maric and diferulic acids.. . . 5. haemodoraceae

1. HANGUANACEAE

Airy Shaw 1965. 1/5 or more. Sri Lanka, Indochina,

Malesia, Palau Islands, Australia.

Hanguana.

Bentham and Hooker (1883), Baillon (1892), Lotsy

(1911), Hutchinson (1934, 1973) and many others

included Hanguana in the Flagellariaceae. However, it

differs markedly from the Flagellariaceae in many

respects (Erdtman 1952; Smithson 1956; Tomlinson

1969). Endlicher (1841) included Hanguana in

Astelieae, which he considered “Genera Juncaceis

afﬁ nia,” and the afﬁ nity to the Asteliaceae has been

accepted by Dahlgren et al. (1985) and Takhtajan

(1987). However, there are many important differences

between Asteliaceae and Hanguanaceae, including tet-

racytic stomata, inaperturate pollen grains, presence of

funicular obturator, bowl-shaped seeds, and peculiar

segmented and starchy endosperm. According to

Stevenson and Loconte (1995), and Rudall, Stevenson

and Linder (1999) the Hanguanaceae are related to the

Zingiberales. According to molecular data (Chase

et al. 2005) Commelinaceae and Hanguanaceae are

sister taxa. Both families contain silica bodies in leaves

and ferulic acid in cell walls (Harris and Hartley 1980;

Rudall and Caddick 1994).

2. COMMELINACEAE

Mirbel 1804 (including Cartonemataceae Pichon

1946, Ephemeraceae Batsch 1802, Tradescantiaceae

Salisbury 1807). 40/650. Tropical and subtropical

regions, especially Africa, southern Asia, Mexico, and

northern Central America with relatively few represen-

tatives in temperate parts of eastern Asia, southern

North America, and Australia. Subfamily Cartone-

matoideae is conﬁ ned to the Southern Hemisphere.