Takhtajan Armen. Flowering Plants

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Superorder ARECANAE 695

syncarpous gynoecium. Fruits 1–3-seeded, with thin

or thick, ﬂ eshy or spongy mesocarp, covered with

reﬂ exed imbricate scales. – eugeissoneae: Eugeissona;

lepidocaryeae: Oncocalamus, Eremospatha,

Laccosperma, Raphia, Lepidocaryum, Mauritia,

Mauritiella; calameae: Korthalsia, Eleiodoxa,

Salacca, Metroxylon, Pigafetta, Plecto comia,

Myrialepis, Plectocomiopsis, Calamus (including

Calospatha), Retispatha, Daemonorops, Ceratolobus,

Pogonotium.

1.2 NYPOIDEAE

Mangrove palm with dichotomously branched,

creeping stem. Leaves paripinnate, reduplicate.

Flowers unisexual, monoecious, with androecium of

only three stamens and apocarpous gynoecium; carpels

large, asymmetrical, have a vascular system different

from that of all other palms. Fruits with ﬁ brous

mesocarp. – Nypa.

1.3 CORYPHOIDEAE

Dwarf or creeping to large fan palms with solitary to

multiple trunks. Leaves palmate or costapalmate, rarely

entire or pinnate, almost always induplicate. Flowers

solitary or in cincinni, bisexual or unisexual, with apocar-

pous or syncarpous gynoecium. Septal glands occur in

syncarpous genera. Fruits usually with ﬂ eshy mesocarp.

– sabaleae: Sabal; cryso phileae: Schippia, Trithrinax,

Zombia, Cocco thrinax, Hemithrinax, Thrinax,

Chelyocarpus, Cryosophila, Itaya; phoeniceae:

Phoenix; livistoneae: Chamaerops, Guihaia,

Trachycarpus, Rhapidophyllum, Maxburretia, Rhapis,

Livistona, Licuala, Johannesteijsmannia, Pholidocarpus,

Pritchardiopsis, Acoelorraphe, Brahea, Colpothrinax,

Copernicia, Pritchardia, Serenoa, Washingtonia;

chunio phoeniceae: Chuniophoenix, Nanno rrhops,

Kerriodoxa; caryoteae: Caryota, Arenga, Wallichia;

corypheae: Corypha; borraseae: Bismarckia,

Satranala, Hyphaene, Medemia, Latania, Lodoicea,

Borassodendron, Borassus.

1.4 CEROXYLOIDEAE

Small to tall. Leaves pinnate or entire and pinnately

ribbed, induplicate. Flowers unisexual (monoecious

or dioecious) or rarely bisexual, with syncarpous

gynoecium. – cyclospatheae: Pseudophoenix;

ceroxyleae: Ceroxylon, Oraniopsis, Juania,

Ravenea; phytelepheae: Phytelephas, Aphandra,

Ammandra.

1.5 ARECOIDEAE

Very small to large, from erect to repent. Leaves pinnate,

pinnately ribbed or bipinnate, induplicate or reduplicate.

Flowers always unisexual, monoecious or dioecious,

sessile and embedded in axis, with syncarpous, often

pseudomonomerous gynoecium. – iriarteeae:

Dictyocaryum, Iriartella, Iriartea, Socratea Wettinia;

chamaedoreeae: Hyophorbe, Wendandiella,

Synechanthus, Chamaedorea, Gaussia; podoco cceae:

Podococcus;

oranieae: Orania; scle rospermeae:

Sclerosperma; roystoneae: Roystonea: rein-

hardtieae: Reinhardtia; coco seae: Beccariophoenix,

Jubaeopsis, Voanioala, Allagoptera (including

Polyandrococos), Attalea, Butia, Cocos, Jubaea,

Lytocaryum, Syagrus, Paraju baea, Acrocomia,

Barcella, Elaeis, Gastrococos, Aiphanes, Bactris,

Desmoncus, Astrocaryum; mani carieae: Manicaria,

euterpeae: Hyospathe, Euterpe, Prestoea,

Neoniholsonia, Oenocarpus: geonomeae:

Pholidostachys, Welﬁ a, Calyptonoma, Calyptrogyne,

Asterogyne, Geonoma; leopold inieae: Leopoldinia;

pelagodoxeae: Som mieria, Pelagodoxa; areceae:

Archontophoenix, Chamebeyronia, Kentiopsis,

Actinokentia, Actinorhytis, Nenga, Pinanga, Areca,

Alloschmidia, Cyphophoenix, Campecarpus,

Basselinia, Cyphosperma, Veillonia, Burretiokentia,

Physokentia, Neoveitchia, Carpoxylon, Satakentia,

Brongniartikentia, Clinosperma, Cypho kentia,

Moratia, Lavoixia, Dypsis, Lemuro phoenix, Masoala,

Marojejya, Calyptrocalyx, Linospadix, Laccospadix,

Howea, Deckenia, Acan thophoenix, Oncosperma,

Tectiphiala, Drymophloeus, Carpentaria, Veitchia,

Balaka, Normanbya, Wodyetia, Ptychosperma,

Ponapea, Adonidia, Solﬁ a, Ptychoc occus, Brassio-

phoenix, Hedyscepe, Rhopalostylis, Verschaffeltia,

Roscheria, Phoenicophorium, Nephrosperma.

Unplaced members of Arecaceae: Alsmithia,

Bentinckia, Clinostigma, Cyrtostachys, Dictyosperma,

Heterospathe, Hydriastele (including Siphokentia,

Gronophyllum, Gulubia), Iguanura, Lepidorrhachis,

Loxococcus, Rhopaloblaste.

Genus dubious: Dransﬁ eldia.

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Superorder ARECANAE 697

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A. Takhtajan, Flowering Plants, 699

A large group of orders derived from the

Liliidae-Lilianae, most likely from the Amaryllidales-

like ancestor. The Commelinidae are characterized by

such synapomorphies as endosperm with copious

starch, cell walls containing UV-ﬂ uorescent com-

pounds ( ferulic and coumaric acids), silicon dioxide

bodies in leaves, stomatal complexes with well differ-

entiated subsidiary cells, epicuticular waxes of the

Strelitzia-type. The commelinids are rather advanced,

mostly with very specialized inﬂ orescences and often

reduced and anemophilous ﬂ owers. They constitute a

monophyletic group, which is supported both by cla-

distic analysis (Dahlgren and Rasmussen 1983) and

rbcL sequence data (Duvall et al. 1993).

Bibliography

Clark WD, BS Gaut, MR Duvall, and MT Clegg. 1993.

Phylogenetic relationships of the Bromeliﬂ orae-

Commeliniﬂ orae-Zingiberiﬂ orae complex of monocots

based on rbcL sequence comparisons. Ann. Missouri Bot.

Gard. 80: 987–998.

Givnish T J, TM Evans, JC Pires, and KJ Sytsma. 1999.

Polyphyly and convergent morphological evolution in

Commelinales and Commelinidae: Evidence from rbcL

sequence data. Molec. Phylogen. Evol. 12: 360–385.

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logenetic trees with distant outgroups: a case study from the

commelinoid monocots. Molec. Biol. Evol. 19: 1769–1781.

Superorder BROMELIANAE

Order 23. BROMELIALES

Mostly short-stemmed epiphytic perennial herbs or

sometimes terrestrial xerophytes. Many species of epi-

phytic genus Tillandsia have elongate stem, and species

of Puya have more or less woody stem, which in

P. raimondii reaches a height of 10 m; in some spe-

cies of Pitcairnia stem is climbing. In epiphytic species

roots often serve as climbing roots. Sacs with bundles

of calcium oxalate raphides occur in all parts as well

as small, spherical silica bodies in the inner periclinal

walls of the epidermal cells of stem and leaves. Sieve-

element plastids of form-P2c. Vessels only in roots, in

roots and stem, or also in leaves, with scalariform or

less often simple perforations. Leaves spirally arranged

or rarely (some species of Tillandsia) distichous,

mostly in basal rosettes, less often cauline, often rigid

and xeromorphous, with broad, sheathing base, simple,

entire or spinose-derrate, linear, broadly subulate, lan-

ceolate, or rarely ovate, parallel-veined, generally with

internal water-storing mesenchymatous tissue and with

characteristic water-absorbing, peltate scales with uni-

seriate stalk, and multicellular, stellate to platelike

shield which is often pressed against the epidermis,

often forms a vase-shaped rosette in which water accu-

mulates and from which it is absorbed. Stomata tetra-

cytic (with two narrow lateral and two short, terminal

subsidiary cells) or sometimes with six subsidiary cells.

Flowers in simple or compound racemes, spikes, heads,

or panicles, rarely solitary, commonly with well-

developed, brightly colored, spathaceous bracts, bisex-

ual or sometimes functionally unisexual, actinomorphic

or rarely somewhat zygomorphic, almost always

3- merous. Perianth segments six in two more or less

unequal cycles; outer segments generally much smaller,

often hyaline or greenish, free or more or less connate;

inner segments petaloid and usually brightly colored,

free or basally connate into short tube, often basally with

ligular appendages. Stamens six in two cycles; ﬁ laments

narrow, inserted at the base of perianth segments when

they are free, and adnate to them when they are connate;

Subclass IV. COMMELINIDAE

700 Subclass IV. COMMELINIDAE

anthers often long and linear, tetrasporangiate, basiﬁ xed

or dorsiﬁ xed, opening longitudinally, introrse. Tapetum

plasmodial. Microsporo genesis successive. Pollen

grains generally 2-celled, 1-colpate, inaperturate or

2-polyporate (restricted to Brome lioideae), subtectate,

mostly reticulate. Gynoecium of three united carpels;

style usually long, slender, more or less 3-lobed, with

different kinds of stigma (conduplicate-spiral or less

often convolute-bladed, simple-erect, cupulate and

coralliform, see Brown and Gilmartin 1984, 1988);

ovary superior to inferior, 3-locular, usually with more

or less numerous ovules. Septal nectaries present.

Ovules anatropous or rarely campylotropous, very

rarely (Hohenbergia) orthotropous, bitegmic, crassi-

nucellate, with parietal cell. Female gametophyte of

Polygonum-type. Endosperm development of a special

subtype of helobial type, in which cell walls are formed

sooner in the chalazal chamber than in the micropylar,

and often the chalazal chamber lacks starch grains

which are copious in the micropylar endosperm

(Dahlgren et al. 1985). Fruits usually septicidal cap-

sules or more often berries. Seeds usually small and

appendaged, rarely naked; seed coat formed by the

inner integument with the various degree of participa-

tion of the outer integument; embryo small or some-

times rather large, cylindrical, mostly lateral, with the

radicle next to the hilum; endosperm copious, mealy,

consisting of large cells containing simple or com-

pound starch grains; peripheral layer of endosperm

represented by smaller cells containing aleurone.

Flavonoles are present. n = 25, rarely 16, 17, 21, 24,

17, 27, 36, chromosomes very small.

Bromeliaceae are set apart from other monocotyle-

dons in many respects. Their synapomorphies include

testal-tegmic seeds (Danilova et al. 1995), unique stel-

late or scalelike multicellular trichomes, solitary silica

bodies, a conspicuously sepaloid and petaloid perianth,

and the unusual conduplicate, spiral stigmas, common

in all three subfamilies (Gilmartin and Brown 1987;

Smith and Till 1998). They are relatively closest to the

Velloziaceae (see Huber 1977; Gilmartin and Brown

1987), and both of them are most probably related to

the Lilianae one hand and to the Commelinanae (espe-

cially Rapateaceae, see Smith 1934) on the other.

Similarities between the Bromeliales and Lilianae

include the presence of raphides, steroidal saponins

and ﬂ avonoids, mostly scalariform perforations of the

root vessels, the presence of septal nectaries, and helo-

bial endosperm. Also, there are deﬁ nite similarities

between the pollen grains of the Bromeliaceae and

those of certain families of the Lilianae, including

Hypoxidaceae (Erdtman and Praglowski 1874) and

Agavaceae (particularly Agave) (Takhtajan 1980).

However, from most of the Lilianae the Bromeliaceae

differ in their stomatal complex (see Tomlinson 1969)

and the presence of silica bodies. In addition, it is

important to mention, that members of the Bromeliales

show UV-ﬂ uorescence in the cell walls, indicating the

presence of the same compounds as are found in the

Arecanae, Commelinanae and Zingiberanae (Harris

and Hartley 1980). Dahlgren et al. (1985) presume that

the ancestors of the Bromelianae belonged to the same

branch as did the ancestors of both Commelinanae and

Zingiberanae (possibly also those of the Arecanae).

Unlike most of the Commelinanae they had retained

the entomophilous syndrome and in this they are still

“liliiﬂ oran”.

1. BROMELIACEAE

A. L. de Jussieu 1789 (including Tillandsiaceae Wilbread

1834). 57/2700 or more. Tropical and partly subtropical

regions of America from Virginia to Patagonia, 1 species

of Pitcairnia in tropical West Africa.

1.1 PITCAIRNIOIDEAE

Nearly always terrestrial. Roots present and largely

functional. Leaves nearly always spinose-serrate.

Ovary superior (or largely so in most genera) to rarely

inferior or semi-inferior (Pitcairnia). Ovules with a

chalazal appendage. Fruits generally septicidal capsules

with an additional loculicidal apical dehiscence in some

genera or rarely indehiscent. Germination epigeal. –

brocchinieae: Brocchinia; pitcairnieae: Navia,

Ayensua, Steyerbromelia, Lindmania, Cotten dorﬁ a,

Connellia, Fosterella, Pepinia, Pitcair nia; puyeae:

Brewcaria, Deuterocohnia, Puya, Encholirium,

Dyckia, Hechtia.

1.2 TILLANDSIOIDEAE

Mainly epiphytic. Roots often functioning as holdfasts,

sometimes lacking. Leaves generally with spineless

margin. Petals appendages present or absent. Ovary supe-

rior or nearly so, rarely (Glomeropitcaimia) semi-inferior.

Ovules with or without chalazal appendages. Fruits

septicidal capsules. – catopsideae: Gatopsis;

glomeropitcairnieae: Glomeropit cairnia; vrueseeae:

Superorder BROMELIANAE 701

Alcanterea, Vriesea, Werauhia; tillandsieae:

Mezobromelia, Racinaea, Guzma nia, Tillandsia,

Viridantha.

1.3 BROMELIOIDEAE

Mainly epiphytic, usually acaulescent. Roots often

functioning as holdfasts. Leaves usually spinose-

serrate or serrulate. Ovary inferior or nearly so. Ovules

usually lack chalazal appendages. Fruits baccate, but

often dry. – Orthophyllum, Cryptanthus, Araeococcus,

Fascicularia, Greigia, Fernseea, Ochagavia, Ursulaea,

Bilbergia, Neoglaziovia, Bromelia, Deinacanthon,

Chevalliera, Pseudananas, Ananas, Acanthostachys,

Hohenbergia, Aechmea, Quesnelia, Canistrum,

Wittrockia, Portea, Pseudaechmea, Hohenbergiopsis,

Androlepis, Lymania, Nidularium, Neoregelia, Strepto-

calyx, Ronnbergia, Disteganthus.

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Superorder ZINGIBERANAE

Order 24. ZINGIBERALES (CANNALES)

Perennial herbs with sympodial, very short to elongate,

often tuberous-thickened and starchy rhizomes or

arboreal with a simple, unbranched trunk lacking sec-

ondary growth (some Strelitziaceae). Silica bodies in

bundle sheath. Vessels only in roots, sometimes pres-

ent also in the shoot, with simple or scalariform perfo-

rations. Leaves distichous or spirally arranged, small

to very large, with an opened or closed sheath, usually

a distinct petiole, and an entire, simple blade, that is

rolled up one side to the other in bud. Stomata para-

cytic or more seldom tetracytic or hexacytic. Flowers

usually in terminal, racemose inﬂ orescences, zygo-

morphic or asymmetrical, bisexual or unisexual. Sepals

three, free or connate to a tube, unlike the petals. Petals

three, sometimes dissimilar, free or connate, some-

times joined to the sepals. Functional stamens one or

ﬁ ve, rarely six (Ravenala and some Ensete), usually one

to several staminodes are present. Anthers basiﬁ xed

or rarely dorsiﬁ xed (some Zingiberaceae), opening

longitudinally. Tapetum secretory or rarely (some

Zingiberaceae and Cannaceae) amoeboid. Microsporo-

genesis successive. Pollen grains usually 2-celled,

inaperturate or more rarely 1-colpate (Zingiber and

Dimerocostus), spiraperturate (Tapeinochilos) and

Superorder ZINGIBERANAE 703

5–6-porate (Costus, Monocostus). Gynoecium of three

united carpels, or one or two of them are more or less

reduced or even obsolete. Ovary superior or semi-

inferior to inferior, with numerous ovules on the intru-

sive placentas. Septal nectaries commonly present.

Ovules anatropous or rarely (some Marantaceae)

campylotropous or (some Zingiberaceae) orthotro-

pous, bitegmic, crassinucellate. Female gametophyte

Polygonum-type or rarely (Costaceae) Adoxa-type.

Endosperm nuclear or rarely (Zingiberaceae and

Costaceae) helobial. Fruits loculicidal capsules, some-

times schizocarpous, nuts or berries. Seeds often oper-

culate next to the radicle, often arillate, seed coat

derived from the outer integument, embryo straight or

curved, endosperm copious with compound starch

grains, or rarely (Maranthaceae) absent, perisperm

usually present, rarely (Strelitziaceae) absent. Contain

raphides and chelidonic acid.

Related to Lilianae and Bromelianae and probably

have a common origin with Bromeliales from ancient

lilealean stock.

Key to Families

1 Functional stamens six or ﬁ ve.

2 The median perianth segment of the inner cycle

not larger than the others. Giant or medium-sized

more or less arborescent plants with erect, woody

in texture trunks or (most species of Strelitzia

except S. nicolai) herbs with short and cormlike

stem, generally with a sympodial starchy rhizome

with branches terminating in aerial shoots. Roots

with anomalous stele ﬁ lled with wide medullary

vessels and scattered phloem islands with a single

sieve tube in each strand. Starch grains ellipsoidal

to spherical or (Ravenala) markedly ﬂ attened.

Silica cells associated with vascular bundles in all

parts of the plant except the roots, silica bodies

druselike or spherical. Raphide sacks present in

the ground tissue of all organs. Vessels in root and

woody stem or (Ravenala) only in roots, with

scalariform and sometimes also simple perfora-

tions. Leaves commonly very large, more or less

petiolate (petiole long to often indistinct or even

lacking), with short basal sheath; lamina with

coarse midrib and closely set parallel, slightly

sigmoid lateral veins interconnected by tertiary

cross-veins and fusing near margin. Stomata

generally tetracytic to hexacytic, often with

sunken guard cells. Flowers in axillary or

(Phenacospermum) terminal thyrse with 1 to sev-

eral lateral, distichously arranged, few-ﬂ owered,

compact, monochasial cymes, each one in the axil

of large, stiff, brightly colored, persistent, spatha-

ceous bract; individual ﬂ owers bi-sexual, more or

less zygomorphic, subtended by carinate bracte-

oles, sweet-smelling. Perianth segments all more

or less petaloid, outer segments free from each

other and more or less equal, the inner ones

slightly or more often strongly dissimilar, the lat-

eral being more or less fused with each other and

in Strelitzia form a stiff, arrow-shaped keel

enclosing functional stamens and style, the

median free or only basally connate with the oth-

ers. Stamens six (Ravenala) or ﬁ ve functional and

one (belonged to the inner cycle) lacking; ﬁ la-

ments elongate, rigid; anthers in Strelitzia with

slightly protruded connective. Style ﬁ liform,

often dilated distally; stigmas 3-lobed, long tur-

binate. Ovary 3-locular, semi-inferior, with con-

spicuous solid extension above the locules.

Ovules numerous in each carpel. Fruits loculi-

cidal capsules. Seeds with brightly colored, hairy

arils, with mostly rudimentary operculum (well

developed only in Ravenala) but without micro-

pylar collar, copious starch- and aleurone-storing

endosperm and 1-layered, starchless perisperm;

embryo straight or curved. Phenylphenalenones

(cyanidin) and ﬂ avonols present (Strelitzia). n =

7, 8, 11. . . . . . . . . . . . . . . . . . 1. strelitziaceae.

2 The median (adaxial) perianth segment of the

inner cycle developed as a large labellum.

Medium-sized or small perennial herbs with

creeping sympodial rhizome rich in compound

starch grains and bearing scalelike leaves and

very short, sometimes branched stem and rather

thick roots. Hat-shaped silica bodies occur in

connection with the vascular bundles, and raphide

sacs occur in all parts. Vessels only in roots, with

scalariform perforations with numerous bars.

Leaves of moderate size, distinctly petioled, with

broadly lanceolate or elliptic lamina with arching

and apically convergent longitudinal lateral veins

emerging in regular intervals from the prominent

midvein and regularly connected by ﬁ ne trans-

versal veinlets; palisade tissue lacking, meso-

phyll of irregularly arranged large and small

cells. Stomata paracytic, with deeply sunken

asymmetrical guard cells. Inﬂ orescences are

704 Subclass IV. COMMELINIDAE

terminal on short leafy branches of the stem; the

partial inﬂ orescences consist of sympodially

connected 1-ﬂ owered monochasial cymes sub-

tended by scalelike bracts; individual ﬂ owers

bisexual, strongly zygomorphic and superﬁ cially

orchid-like, bracteate, malodorous. Perianth seg-

ments six in two cycles, outer segments subequal,

narrow, basally connate into tube with linear-

lanceolate and pointed lobes, inner segments

strongly dissimilar, the median (adaxial) one

large and transformed into variously shaped

labellum, the two lateral ones smaller and over-

lap the stamens. Stamens ﬁ ve, the median (adax-

ial) one of the inner cycle (opposite the labellum)

missing or sometimes staminodial; ﬁ laments

short; anthers with shortly subulate connective

appendage. Style erect, slender, apically widens

into 3-lobed, asymmetric, ﬂ attened, marginally

laciniate-ﬁ mbriate stigmatic lobes. Stigma mono-

symmetric, dorsiventrally ﬂ attened, the base part

of adaxial side contain secretory tissue (viscid-

ium). Ovary has a solid apical extension, and with

numerous ovules per locule. Fruits many-seeded

elongate capsules with papery pericarp termi-

nated by a beak formed by the proximal part of

the ovary extension. Seeds globose to pyriform,

smooth or hairy, with a large lacerate aril and an

operculum in front of the radicle. Testa vascular-

ized, exotesta and next two layers ligniﬁ ed,

endotesta of radially elongated sclereids; testa

encrusted with phytomelan; perisperm slight

(Wen et al. 1997). Embryo cylindrical, endosperm

copious, with isodiametric starch grains, mostly

only one cell layer thick and at the chalazal part

several cell layers thick. n = 9. . . .2. lowiaceae.

3 Flowers bisexual. Commonly very tall herbs

with creeping sympodial rhizomes, nearly

acaulescent or with slender, unbranched aerial

stems. Roots with normal polyarch stele or

with occasional internal phloem strands.

Starch grains cylindrical or ellipsoidal but

never ﬂ attened. Laticifers lacking. Silica cells

( stigmata) internal, adjacent to vascular bun-

dles, with unequally thickened walls, each

including a small rectangular silica body, each

with a deep central depression. Vessels only in

roots, with scalariform perforations. Leaves

distichous, with long, basal sheath and petiole

(sometimes indistinct or lacking). Stomata tet-

racytic or less often paracytic, with narrow

deep subsidiary cells adjacent to guard cells.

Flowers in terminal ﬂ attened thyrse consisting

of distichously arranged, dense, monochasial

cymes (cincinni), each one in the axil of large

or rather small, usually brightly colored,

keeled or more often boat-shaped bract; indi-

vidual ﬂ owers subtended by carinate bracte-

oles, strongly zygomorphic. Perianth segments

all petaloid, dissimilar, all fused mutually and

with the ﬁ lament bases to form a basal tube;

above this the median posterior (adaxial) one

of the outer cycle and the ﬁ laments are dis-

tinct, whereas the other ﬁ ve segments are fur-

ther connate into a 5-dented or 5-lobed,

boat-shaped upper lip. Functional stamens

ﬁ ve, with the ﬁ laments adnate to the tubular

part of the perianth, the outer median stamen

staminodial, inserted on the top of the perianth

tube opposite the median sepal. Ovules soli-

tary in each locule. Fruits schizocarps with a

rather thin pericarp splitting into (2)3 usually

blue, ﬂ eshy mericarps each of which contains

a single seed. Seeds exarillate, with an opercu-

lum formed by the tissue of pericarp, whereas

only the rudimentary operculum if any are

formed by the outer integument (Plisko 1985);

perisperm represented by thin aleurone-stor-

ing layer, but endosperm copious, starch- and

oil-storing; embryo straight or curved. n = 11,

12. . . . . . . . . . . . . . . . . . . .4. heliconiaceae.

3 Flowers unisexual. Large to gigantic herbs with

short, massive underground rhizome. Root stele

anomalous in possessing numerous scattered

vessels and phloem islands, each phloem strand

with several wide sieve tubes. Starch grains

ﬂ attened and eccentric. Articulated laticifers

occur in association with vascular bundles in all

parts except roots. Silica present in all parts

except roots as small, more or less rectangular,

troughlike bodies in cells (stigmata) with

unevenly thickened walls. Vessels in roots, with

scalariform and simple perforations. Leaves

spirally arranged, with well-developed, coarse,

tubular sheath and easily frayed lamina with

coarse, compound midvein and closely set, par-

allel, slightly sigmoid lateral veins intercon-

nects by tertiary cross-veins and fusing near

margin. Stomata tetracytic. Flowers in terminal

Superorder ZINGIBERANAE 705

thyrse which consists of spirally arranged lat-

eral dense and compact monochasial cymes

(cincinni) each in the axil of large, ﬁ rm, keeled

or boat-shaped bracts; individual ﬂ owers are

subtended by hyaline, recurved bracts and uni-

sexual, either female (basally in thyrse), with

nonfunctional stamens, or male (apically in

thyrse), with nonfunctional gynoecium.

Perianth segments petaloid, dissimilar, the three

segments of outer cycle and two segments of

the inner cycle united into usually 5-dented or

5-lobed, initially tubular structure that soon

splits on one side, the third (adaxial) segment of

the inner cycle is distinct from the others and

directed downward. Functional stamens 6 (most

species of Ensete) or the median (adaxial) of

the inner cycle missing. Ovary 3-locular, inferior;

ovules 10–50 in each locule. Fruits berries.

Seeds exarillate, with well-developed opercu-

lum and micropylar collar; peri-sperm repre-

sented by thin layer, but endosperm copious,

starchy and mealy; embryo straight or slightly

curved (Ensete). Alkaloids (indole), and ﬂ a-

vonols (kaempferol and quercetin) present or

absent, phenylphenalenones (cyanidin, or cya-

nidin and delphinidin) present; n = 7, 9–14,

16, 17. . . . . . . . . . . . . . . . . . . . . 3. musaceae.

1 Functional stamen one.

4 Leaves distichous. Aromatic herbs. Cells con-

taining ethereal oils present in the parenchyma

of all parts of the plant and very common in the

abaxial leaf epidermis. Tannin cells in ground

parenchyma are common in all parts. Silica

commonly restricted to epidermis of lamina,

either as silica sand or as solitary, irregularly

spherical bodies. Starch grains often ﬂ attened.

Vessels chieﬂ y in roots and stems, with scalari-

form, or occasionally with simple perforations.

Leaves have long open or rarely closed sheath,

with or without ligule. Two sterile lateral sta-

mens of the inner cycle united to form the peta-

loid labellum: two lateral staminodia of the outer

cycle petaloid or inconspicuous, ﬂ anking the

stamen or adnate to the labellum: the median

anterior member of the outer cycle always

reduced; anthers usually opening longitudinally,

occasionally dehiscing by pores. Pollen grains

inaperturate or rarely (Zingiber) 1-colpate; exine

very thin, hardly resistant to acetolysis, but the

intine well developed and lamellated. Septal

nectaries (2, sometimes 3 or 1) of various shape

present at the style base. Ovary at ﬁ rst always

3-locular but when fully developed either incom-

pletely 3-locular or 1-locular and with parietal

or basal-parietal placentation; ovules numerous

to many, anatropous or orthotropous. The chala-

zal chamber of the endosperm degenerates quite

early, often as early as the two-celled stage.

Fruits mostly loculicidally dehiscing capsules,

sometimes indehiscent or ﬂ eshy, berrylike.

Seeds with thin, generally lobed or laciniate aril

(except Caulokaempferia coenobiales) and are

ﬁ lled up by starchy perisperm (with mostly com-

pound starch grains) surrounding the thinner

endosperm that may contain aleurone grains and

sometimes also starch grains; chalazosperm

poorly developed or absent; embryo straight

(except in Alpinia), usually reaching the exten-

sion of the perisperm, rarely shorter; testa

encrusted with phytomelan. Phenylpropanoids

and related curcumins, ethereal oils present. n =

8–21 and 25, mostly 12. . . . 5. zingiberaceae.

4 Leaves spirally or four-ranked arranged. Non-

aromatic herbs with multicellular hairs. Oil and

tannin cells lacking. Silica bodies stellate, druse-

shaped, restricted to internal cells next to vascu-

lar bundles. Starch grains not ﬂ attened, often

large and irregular. Vessels mainly in roots and

stems, usually only with scalariform perfora-

tions. Leaves with short, closed tubular sheath;

ligule present. Stomata paracytic or tetracytic.

Flowers solitary or in terminal or axillary inﬂ o-

rescences, bracteate and sometimes bracteolate.

Perianth segments with distinct calyx and

corolla; calyx imbricate Fertile stamen often

petaloid. All staminodia (2 of the inner cycle

and 3 of the outer one) fused into a conspicuous

labellum which is united with the basal part of

the stamen into a papillate tube. Pollen grains

very large, distinctly aperturate, 1–2-colpate,

spiraperturate or 5–16-porate, resistant to ace-

tolysis. Ovary 3-locular (Costus) or 1–2-locular;

ovules numerous per locule, anatropous The

chalazal chamber of the endosperm is coeno-

cytic and remains at the time when the embryo is

differentiated. Septal nectaries two, immersed in

the apical part of the ovary. Fruits capsules

dehiscing loculicidally or rarely explosively or