Takhtajan Armen. Flowering Plants

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Superorder PANDANANAE 685

3-lobed, capitate or clavate stigma; ovary variously

pubescent on the outside, inferior or rarely

semi-inferior (almost superior in Vellozia burle-

marxii), 3-locular, with numerous ovules on stalked

intrusive placentas. Ovules anatropous, bitegmic,

weakly crassinucellate to tenuinucellate, with funicu-

lar obturator, without parietal cell. Female gameto-

phyte of Polygonum-type. Endosperm helobial or

(Acanthochlamys) nuclear (Li and Gao 1993). Fruits

capsular, loculicidal or opening by pores, often ﬂ at or

concave on the top, crowned with the scar of the peri-

anth, or 6-toothed, sometimes spiny, with numerous

seeds. Seeds rather small, black; seed coat testal-

tegmic, formed by both integuments, without phy-

tomelan but with phlobaphene in exotesta; endosperm

copious, in its outer layers containing aleurone and

lipids and in the inner layers plenty of globose starch

grains; embryo small, cylindrical or narrowly ovoid,

n = 7, 8, 17, 19, 24.

Possibly related to the Bromeliaceae (Huber 1969,

1977; Dahlgren et al. 1985; Takhtajan 1987), with

which Velloziaceae share such common characters, as

the para- and tetracytic stomata, helobial endosperm

formation, testal-tegmic seed coat, and starchy

endosperm. Afﬁ nity with the Bromeliaceae is also

supported by the presence of the same kind of epicu-

ticular waxes (Frölich and Barthlott 1988), but the lack

of the UV ﬂ uorescence of the cell walls is at variance

with this allocation (Kubitzki 1998). As regards the

placement of the family together with the Cyclantha-

ceae, Pandanaceae, Stemonaceae and Acanthochlamys

in the rbcL trees (Chase et al. 1995), I agree with

Kubitzki (1998) that it “does not seem to be accompa-

nied by morphological characters: at best it indicates

the strong isolation of the family”.

Key to Families

1 Anthers tetrasporangiate. Perennial herbs and

sparsely branched shrubs. Stems are covered with

persistent imbricate sheaths of fallen leaves on the

upper part and on the persistent adventitious roots,

piercing the withered leaf bases on the lower part.

Raphides sometimes present. Leaves in a basal

rosette or apical tufts, sometimes up to 1 m long,

simple, linear-acuminate, with midrib, sometimes

with a saw-toothed margin. Flowers solitary to

12–15 on each scape, pedicellate, terminal, bisex-

ual, rarely functionally unisexual and the plants

dioecious (Barbaceniopsis), actinomorphic, often

large. Perianth segments six in two whorls, free or

basally connate, sometimes (Barbacenia) with a

small corona behind the anthers. Stamens 6–48,

up to 76 (in Vellozia) in six bundles when more

than six, lacerated staminal appendages often pres-

ent in Vellozia; ﬁ laments epigynous (Talbotia) or

adnate to the perianth, sometimes ﬂ attened; anthers

long; pollen grains 1-sulcate or nonaperturate.

Septal nectaries conspicuous. Ovary inferior or

rarely semi-inferior, stigma large, erect or spread-

ing. Fruits loculicidal capsules, or dehiscent by api-

cal, basal, or longitudinal slits. Seed small,

numerous; exotesta thickened or not (tegmen tan-

niniferous – Pleurostima); embryo small, endosperm

copious, hard, with hemicellulose, protein, oil and

starch. Biﬂ avonoids present. n = 7, 8, 17, 24 (x =

12?).. . . . . . . . . . . . . . . . . . . . . 1. velloziaceae.

1 Anthers disporangiate. Perennial herbs with short

rhizome and thin, long ﬁ briform roots. Stems

without mantle of adventitious roots. Raphides

and tannin cells absent. Sieve-element plastids

form-P2c. Vessels with simple perforations and

helical thickenings present in the root, scape and

leaves. Leaves acerose, dorsiventral, ventrally

subsemiorbicular and 2-canaliculate, dorsally ﬂ at-

tened and 1-canaliculate, sheathed at the base.

Midrib is composed of two “back-to-back”

vascular bundles. Stomata paracytic. Inﬂ orescence

a compound capitulum on a scape arising from the

rhizome, at the base usually surrounded by three

leaﬂ ike aristate bracts, the peduncle bearing 5–8

few-ﬂ owered capityla, the ﬂ owers subtended by

aristate bractlets. Flowers bisexual, actinomor-

phic, shortly pedicellate, perianth segments six in

two cycles, connate into tube. Stamens six in two

cycles, borne upon the corolla lobes; ﬁ laments

short; anthers oblong, dorsiﬁ xed, dehiscent by

longitudinal slits. Pollen grains 2- or 3-nucleate,

spheroidal, 1-colpate, ﬁ nely verrucate to reticu-

late. Septal nectaries absent. Fruits trigonous cap-

sules. Seeds oblong, brown, tegmic, but the

mechanical strength is provided mainly be the

thickened periclinal walls of the cell layers imme-

diately beneath the seed coat; this layer contains

aleurone and free of starch; the rest of the

endosperm contains starch in the form of com-

pound starch grains; embryo large (Kao 1989; Kao

and Kubitzki 1998). Steroid saponins may be pres-

ent. n = 19.. . . . . . . 2. acanthochlamydaceae.

686 Subclass II. LILIIDAE

1. VELLOZIACEAE

Hooker 1827 (including Barbaceniaceae Arnott 1842).

8/250. America from Panama to Argentina (especially

in southeastern Brazil), Madagascar, tropical and

southern Africa, southwestern Arabia.

Talbotia, Xerophyta, Barbaceniopsis, Barbacenia

(including Aylthonia), Pleurostima, Burlemarxia,

Nanuza, Vellozia.

2. ACANTHOCHLAMYDACEAE

P.C. Kao 1989. 1/1. Subalpine xerophytic valley zone

of southeastern Tibet, southwestern China and western

Sichuan.

Acanthochlamys

According to Kao and Kubitzki (1998), a possible

relationship of the Acanthochlamydaceae should be con-

sidered with all those families that share with them the

possession of epigyny and starch accumulation in the

endosperm, viz. Bromeliaceae, Haemodoraceae and

Velloziaceae. They ﬁ nd it signiﬁ cant that these three fam-

ilies agree with Acanthochlamys in having starch grains

composed of relatively few elements in the endosperm.

Among these families, the Bromelia ceae differ too much

from Acanthochlamys in their perianth differentiation

into calyx and corolla and their epidermal cells contain-

ing silica bodies. The remaining two families agree with

Acanthochlamys in the collapsed testa and 2-layered teg-

men, the Velloziaceae also in the disporangiate anthers,

and additionally the Haemodoraceae in the thickened

outer periclinal wall of the aleurone layer. However, the

absence of cell wall-bound ferulate is shared only with

the Velloziaceae. Therefore, Kao and Kubitzki favour the

afﬁ nity with the Velloziaceae, although in their opinion

the relationship “may not be very close”. According to

Behnke et al., their “combined results, i.e. small form-P2c

plastids without loosely packed crystals, clear distance in

Neighbor Joining and strong bootstrap support, favour a

monotypic family whose closest afﬁ nities are with

Velloziaceae, and with no close afﬁ nities to

Amaryllidaceae” (2000: 117).

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Kao PC. 1989. Acanthochlamydaceae – a new monocotyledon

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483–507.

Superorder PANDANANAE 687

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ology of Acanthochlamys bracteata: morphological and ana-

tomic studies on vegetative organs. J. Sichuan Univ. 32:

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fertilization, embryogenesis and endosperm development of

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30: 260–263 (in Chinese with English summary).

Li P, P-C Gao, F Chen, and HX Luo. 1992. Studies on morphol-

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Ecol. 19: 483–495.

Order 20. STEMONALES

Perennial rhizomatous herbs with erect stems or trail-

ing or twining herbs with fasciculate tuber. Raphides

present or absent. Vessels only in roots or also in stem

and even leaves, with scalariform perforations. Leaves

alternate, opposite or verticillate, petiolate or sessile,

lanceolate to ovate, with parallel, arcuate-striate or

pinnate-striate venation. Stomata anomocytic or

(Pentastemonaceae) tetracytic. Flowers solitary (axil-

lary or terminal) or in few-ﬂ owered cymes or racemes,

bisexual or rarely unisexual, actinomorphic, 2-merous,

3–4-merous, or rarely (Pentastemona) 5-merous.

Perianth segments free or sometimes more or less con-

nate, similar or more or less dissimilar. Stamens

isomerous with the perianth segments; ﬁ laments free

or basally connate, elongate or short; anthers tetraspo-

rangiate, basiﬁ xed, opening longitudinally, introrse,

688 Subclass II. LILIIDAE

often produced in sometimes rather long appendages.

Tapetum secretory. Microsporogenesis successive.

Pollen grains 1-colpate or (Pentastemona and

Stichoneuron) inaperturate, reticulate, spiny-gammate

or psilate (Furness and Rudal 2006). Gynoecium of

two or 3–6(-8) or more united carpels; stylodia free or

more or less connate; ovary superior to inferior, 3–6(-8)-

locular or 1-locular, with various types of placentation.

Ovules few or numerous, anatropous or hemitropous,

bitegmic, crassinucellate, with parietal placentation.

Female gametophyte or Polygonum-type. Endosperm

nuclear. Fruits capsules (sometimes ﬂ eshy and berry-

like) or less often berries. Seeds ellipsoidal to globose,

usually with various kind of appendages; seed coat

derived mainly from testa; endosperm copious, con-

tains aleurone, fat, and starch grains; embryo usually

small and nearly undifferentiated. Subterranean parts

of Stemona and Croomia contain insecticidal alkaloids

known only from these genera.

Key to Families

1 Ovary superior or semi-inferior. Flowers 2-merous.

Twining or trailing, glabrous herbs with short rhi-

zome and often tuberous roots or erect herbs with

horizontally creeping rhizome and with scale leaves.

Stems vascular bundles in one or two rings. Leaves

alternate, opposite or verticillate, simple, linear, lan-

ceolate, ovate, or triangular, petiolate or subsessile,

cuneate to cordate or hastate at base, acuminate;

venation parallel to arcuate-striate or arcuate-pin-

nate, with parallel transverse veinlets. Flowers large

to rather small, in axillary few-ﬂ owered cymes or

solitary, bisexual or rarely (Stichoneuron) unisex-

ual, articulated from pedicel. Perianth segments

four in two cycles, acuminate, free, sepaloid or pet-

aloid. Stamens four in two cycles; ﬁ laments short,

inserted at base of perianth; anthers in Stemona

with appendaged connective. Pollen grains 1-col-

pate, tectate-columellate to granular, 1-colpate or

(Stichoneuron) inaperturate, with microretivulate or

reticulate (Croomia) ornamentation. Gynoecium of

two carpels; stigma sessile; ovary semiinferior,

1-locular, with few to many, basal (Stemona) or api-

cal ovules. Fruits 2-valved capsules. Seeds longitu-

dinally ridged, with elaiosome of juicy, uniseriate or

vesicular hairs from hilum raphe or micropyle; testa

several-layered, ridges many cells high, tanninifer-

ous; endosperm stores aleurone, lipids and some

starch. n = 9, 12. . . . . . . . . . . . . . .1. stemonaceae.

1 Ovary inferior, ﬂ owers 5-merous. Small, succulent

monopodial herbs with scale leaves on short vertical

rhizome; hairs uniseriate. Stem prostrate or slightly

ascending, unbranched, ﬂ eshy, juicy, loosely attached

to substratum with slender adventitious roots. Leaves

alternate, dispersed, long-petiolate, shortly sheathing

at the base, cordate or rounded at base, acute, sub-

acuminate, with arcuate- pinnate venation and trans-

verse parallel veinlets; petioles and sheathing leaf

bases ciliate. Stomata paracytic, or tetracytic to ency-

clocytic. Flowers small, in axillary, simple or

compound racemes, wholly or partly functionally

unisexual, bracteate and with large dorsal bracteole;

pedicels not articulated. Perianth segments ﬁ ve, in

one cycle, almost free or partially connate into long

to short tube, with ﬁ ve rounded imbricate lobes.

Stamens ﬁ ve, in one cycle, attached to base of peri-

anth tube; ﬁ laments very short, united into short,

ﬂ eshy ring; anthers subsessile; connective broad,

their extensions contacting stigmas. Pollen grains

intectate, inaperturate, scabrate. Gynoecium of three

carpels; style short and thick, with ﬁ ve apparently

nectariferous pouches at its base inside staminal

tube; ovary inferior, provided with ten longitudinal

ﬂ anges or ribs, 1-locular, with numerous anatropous

ovules on intrusive placentas. Fruits berries, dis-

tinctly ribbed, containing many seeds. Seeds oblong,

endotestal. with 10–13 faint longitudinal ridges,

provided with inﬂ ated funicular arillode covering

about one third of the seed, with sarcotestalike hya-

line exotesta; endotesta with massive U-shaped

thickenings; endosperm copious, contains aleurone

and lipids as well as starch grains mainly around

embryo and near chalaza; embryo minute. n = 7.

. . . . . . . . . . . . . . . . . . . . . . . 2. pentastemonaceae.

1. STEMONACEAE

Caruel 1878 (including Roxburghiaceae Wallich 1832;

Croomiaceae Nakai 1937). 3/30. From India, southern

China, and Japan through Malesia to Australia and

southeastern United States (one species of Croomia).

Stemona, Croomia, Stichoneuron.

2. PENTASTEMONACEAE

Duyfjes 1992. 1/2. Sumatra.

Superorder DIOSCOREANAE 689

Pentastemona.

Related to the Stemonaceae, but markedly differ in

5-merous perianth and androecium, inferior ovary,

prominent stigmatic lobes, parietal placentation, ber-

rylike fruits, several-layered testa, intectate pollen

grains, alternate leaves, leaf venation, compound mid-

rib, tetracytic or encyclocytic stomata.

Bibliography

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Superorder DIOSCOREANAE

Order 21. DIOSCOREALES

Perennial herbs with twining or trailing or less often

erect shoots arising from a ﬂ eshy, starchy rhizome or a

tuber arising by hypertrophy of the internodes above the

cotyledon (epicotyl) or/and by a lateral hypertrophy of

the hypocotyl; the tuber with or without secondary

growth. Roots commonly mycorrhizal, without root

hairs. Tannin cells mostly present or (Tricho podaceae)

lacking. Raphides in mucilaginous idioblasts generally

present. Vessels in roots and sometimes also in stem,

with scalariform perforations. Sieve-element plastids of

P2c- or P1/2c-type (Behnke 2002). Leaves all basal,

alternate, opposite or verticillate, sessile or more often

petiolate, entire to compound, with various kinds of

venation; petiole mostly with a pulvinus at each end.

Ptyxis conduplicate or (Taccaceae) plicate. Stomata

anomocytic. Flowers in various kind of inﬂ orescences

or solitary, bisexual or more often unisexual, actinomor-

phic, 2-merous, 3-merous. Perianth segments four, six

in two cycles, more or less similar, usually basally con-

nate into a short tube. Stamens isomerous with peri-

anth segments; inner ones sometimes staminodial or

obsolete; ﬁ laments free or adnate to the perianth seg-

ments; anthers basiﬁ xed, tetrasporangiate, introrse or

sometimes extrorse, opening longitudinally, often with

apically projected connective. Tapetum secretory.

Microsporo genesis simultaneous or (Trichopus) suc-

cessive. Pollen grains 2-celled, 1-, 2- or 4-colpate (sul-

cate) or (Avetra) 4(5)-porate and spinulose. Gynoecium

of three united carpels; stylodia free or connate; ovary

superior, semi-inferior or mostly inferior, 1- or 3-locu-

lar, with 1–2 or numerous ovules per locule. Ovules

anatropous, bitegmic, crassinucellate or (Trichopodaceae)

tenuinucellate. Female gametophyte of Polygonum-type

690 Subclass II. LILIIDAE

or very rarely (in some species of Dioscorea) tetrasporic

of Drusa-type. Endosperm nuclear. Fruits mostly cap-

sules, rarely indehiscent and samaroid (Trichopus and

Rajania) or berries. Seeds of various shapes, often

(Dioscorea and Stenomeris) winged; seed coat formed

by both the integuments (endotestal and exotegmic),

lack phytomelan but often, especially those in capsular

fruits, contain phlobaphene; endosperm copious, con-

tains aleurone, lipids, some starch, and commonly also

hemicellulose in form of cell-wall thickening (absent in

Stenomeris); embryo small, but well differentiated, with

subterminal plumule and broad, ﬂ at lateral cotyledon, or

minute, undifferentiated. Often produce steroidal

saponins, chelidonic acid, and sometimes (species of

Dioscorea) alkaloid dioscorine. Flavones apparently

lacking, although ﬂ avone C-glycosides may be present

in the Trichopodaceae (Williams et al. 1988).

Closely related to the Stemonales and have a com-

mon origin with them.

Key to Families

1 Ptyxis plicate. Embryo minute, undifferentiated,

basal. Acaulescent perennial herbs from starchy,

more or less tuberous-thickened, globose or elongate

rhizome containing alkaloids. Raphide cells present

in stem and leaves. Vessels only in roots, with sca-

lariform perforations. Leaves all basal, with basally

more or less widened and often long petioles and

entire to much dissected laminas; venation arcuate-

striate, palmate-striate, or pinnate-striate with reticu-

late veinlets. Midrib compound. On both sides of

leaves occur peculiar trichomes consisting of a short

row of cells bearing a multicellular body on which is

another cell row at the apex. Mesophyll with scat-

tered mucilage cells. Stomata anomocytic or sur-

rounded by one cell. Inﬂ orescences umbellate,

apparently cymose, on long peduncle, the bracts

forming an involucre, the inner ones narrower and

often ﬁ liform; ﬂ owers bisexual, actinomorphic,

3-merous. Perianth segments six in two cycles,

imbricate, with a short, broad tube and six lobes,

these biseriate, somewhat petaloid, the inner ones

usually longer than the others. Stamens six in two

cycles; ﬁ laments short, inserted on perianth tube,

with inﬂ exed margins, apically forming, together

with the broad connectives, hoods over anthers;

anthers tetrasporangiate, opening longitudinally,

introrse. Pollen grains 2-celled, 1-colpate,

tectate-columellate, verrucate to striate. Gynoecium

of three united carpels; style short, with three broad

stigmas often petaloid and reﬂ exed over the style;

ovary 6-ribbed, inferior, 1-locular, with numerous

pendulous ovules on more or less intruded parietal

placentas. Septal nectaries present in several spe-

cies. Ovules anatropous, apotropous, with parietal

cell. Fruits with a ﬂ eshy pericarp, 6-ribbed, berry-

like, irregularly disintegrating, rarely loculicidal

capsules (Tacca subgenus Schizocapsa). Seeds of

various shapes, mostly provided with longitudinal

ridges, sometimes with thin ﬂ eshy aril and ﬂ eshy

raphe; seed coat formed by both the integuments

(both 2-layered as in Pentastemona) which are well

developed and take part in the mechanical function

(restricted in the endotesta and exotegmen) and

contains phlobaphene; endosperm copious, con-

tains aleurone and lipids, but not starch; embryo

minute, basal, undifferentiated, globose or ovoid,

with laterally inserted cotyledon (as in Dioscorea).

n = 15. . . . . . . . . . . . . . . . . . . . . . . . 1. taccaceae.

1 Ptyxis conduplicate. Embryo small, but well differ-

entiated, with subterminal plumule and broad ﬂ at

cotyledons.

2 Flowers bisexual. The connective produced into

a long appendage.

3 Fruits many-seeded loculicidal capsules. Tall

climbers with short, subterranean horizontal

rhizome. Stem tough, twining to the left.

Leaves alternate, cordate, acuminate, arcuate-

striate veined with many primary veins con-

nected by more or less transverse commissural

veinlets. Flowers in lax axillary panicles or

solitary, pedicillate, pedicels bracteolate.

Perianth segments connate into urn-shaped,

6-lobed tube. Stamens six, inserted in one

series near the mouth of tube; ﬁ laments short,

deﬂ exed; anthers with broad connective that is

apically prolonged into an elongated append-

age with spathulate, horned apex. Style short,

columnar, ending in three biﬁ d stigmas; ovary

with many ovules in each locule. Capsules lin-

ear and much elongated, 3-winged, with thin

pericarp. Seeds ﬂ at, triangular, with large

membranous wing on one side at the top; cells

of mesotesta intensively coloured by phloba-

phenes; cells of endosperm thin-walled.. . . . .

2. stenomeridaceae.

Superorder DIOSCOREANAE 691

3 Fruits indehiscent or opening by irregular rup-

tures, dry or slightly ﬂ eshy, 1- to 6-seeded.

Erect herbs with short rhizome. Leaves entire,

petiolate, ovate- elliptic to cordate-sagittate,

3–7-veined; primary veins palmate, secondary

venation reticulate. Flowers solitary and axil-

lary or in terminal or axillary short racemes or

fascicles, with long pedicels. Perianth more or

less campanulate. Stamens six, inserted at

base of perianth segments; ﬁ laments very

short; anthers with broad connective that is

apically prolonged into long pointed append-

age, introrse. Pollen grains 1-colpate, bicon-

vex and not spinulose (Trichopus) or

4(5)-porate, spheroidal, more or less spinulate

(Avetra). Style short, columnar, with 3-lobed

stigma, the lobes biﬁ d; ovary with two ovules

per locule. Ovules tenuinucellate (at least in

Trichopus). Fruits three wings or keels. Seeds

more or less compressed, with deeply fur-

rowed seed coat derived from both testa and

tegmen; exotegmic cells elongated and differ-

entiated as a mechanical tissue; endosperm

copious, strongly ruminate; the outer wall lay-

ers are hard and thick owing to heavy deposi-

tion of hemicellulose; embryo minute, linear,

straight. n = 14.. . . . . . . 4. trichopodaceae.

2 Flowers unisexual, dioecious. Mostly lianous

herbs or rarely shrubby plants with usually tuber-

ous rhizome or woody rootstock that sometimes

have secondary growth, rarely rhizome. Aerial

stem usually climbing or trailing. Leaves alternate

or rarely opposite, entire or lobed and often cor-

date or palmately compound; venation arcuate-

striate or palmate-striate with 3–13 primary veins

and reticulate veinlets; bulbils of cauline nature

occur in leaf axils of many species of Dioscorea.

Flowers small, in spikes, racemes, cymes, or pan-

icles, pedicellate or sessile, often ebracteolate.

Perianth segments usually basally connate into

short tube. Nectaries commonly present. Fertile

stamens six or three, very rarely one, erect or

spreading; ﬁ laments free or sometimes shortly

connate, attached to perianth tube; anthers not

appendaged, rarely apiculate, dorsiﬁ xed, introrse

or sometimes extrorse. Pollen grains 1-colpate,

2-colpate or occasionally 3-colpate. Pistillodia

present or absent. Stylodia free or connate and

apically 3-lobed or 3-brachiate; ovary with two to

many ovules per locule. Staminodia often present

in female ﬂ owers. Fruits usually loculicidal cap-

sules, generally 3-angular or 3-winged, rarely

samaras (Rajania) or berries (Tamus). Seeds

winged or not, ﬂ at or scarcely compressed, rarely

(Tamus) globose, not ribbed. Endotesta is mostly

crystalliferous and in each cell a single crystal of

calcium oxalate is embedded in a moiety of

phlobaphene and sclerotic exotegmen consists of

cells with pitted or sculptured walls (Huber 1998).

n = 8, 9, 10, 12.. . . . . . . . . . . . 3. dioscoreaceae.

1. TACCACEAE

Dumortier 1829. 1/13 (or 35–50 ?). Pantropical, but

best developed in Southeast Asia and Polynesia.

Tacca.

2. STENOMERIDACEAE

J. Agardh 1858. 1/2. Peninsular Malaya, northern

Borneo and the Philippines.

Stenomeris.

The most archaic member of the order.

3. DIOSCOREACEAE

R. Brown 1810 (including Tamaceae Martynov 1820;

Tamnaceae J. Kickx f. 1826). 6/650. Pantropical, a few

species in subtropical and warm-temperate regions;

Tamus (4–5), the only genus with baccate fruits, dis-

tributed in Macaronesia, Europe, Mediterranean

region, and western Asia.

Dioscorea, Borderea, Testudinaria, Epipetrum,

Rajania, Tamus.

Closely related to the Stenomeridaceae.

4. TRICHOPODACEAE

Hutchinson 1934 (including Avetraceae Takhtajan

1997). 2/2. India (southwestern Deccan Peninsula), Sri

Lanka, Malay Peninsula (Trichopus) and eastern

Madagascar (Avetra).

Trichopus, Avetra.

692 Subclass II. LILIIDAE

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A. Takhtajan, Flowering Plants, 693

The subclass Arecidae is one of the most ancient phyla

of the liliopsids and probably had a common origin

with the Liliidae. Both subclasses derived from a hypo-

thetical extinct common ancestor, which most likely

was a terrestrial rhizomatous perennial herb of more or

less wet habitats without vessels and with primitive

1-colpate, boat-shaped, pollen grains (2-celled when

shed) and primitive free, conduplicate carpels with

decurrent stigmas. In both subclasses there are still

members with a number of primitive characters,

including apocarpous gynoecium and primitive car-

pels. During the evolution of the Arecidae a progres-

sive reduction and specialization of ﬂ owers has taken

place, which has been compensated by the origin of

very specialized compound inﬂ orescences subtended

by a usually well-developed bract, the spathe.

Superorder ARECANAE

Order 22. ARECALES

Usually large, sometimes very large, rarely rather small

(only 2 cm in diameter and less than 25 cm tall in some

species of Chamaedorea), more or less treelike or

shrubby plants, branched or more often unbranched,

erect or sometimes climbing, rarely stemless, some-

times with creeping rhizomes, but never producing

bulbs or corms; lacking cambium but sometimes

increasing in diameter by primary growth. Silica bodies

common. Calcium oxalate usually occurs as raphides,

rarely as isolated crystals or as crystal sand. Vessels

usually in all organs, in roots usually with simple per-

forations, in stem and leaves mostly with scalariform

perforations (always scalariform in Nypa). Leaves

spirally arranged, usually in terminal rosettes, scattered

along the stem in climbing species, rarely basal, often

large or very large, usually clearly differentiated into

petiole and lamina, with broad, clasping sheath at base;

lamina usually palmately or pinnately divided, less

often entire or bipinnate (Caryota), undivided and

densely plicate in bud, with a stout central axis (costa

or rachis), which is short or absent in truly palmate

leaves, better developed in costapalmate leaves, and

prominent in the pinnate leaves; some laminas are

divided so that the segments (of palmate leaves) and

pinnae (of pinnate leaves) appear to be either V-shaped

(termed induplicate) while the others are so divided

that they appear to be like an inverted V (termed redu-

plicate). Stomata tetracytic. Epicuticular wax of

Strelitzia type. Flowers usually small, numerous, in

usually large, compound or rarely simple spikes or

panicles, sometimes in dichasia, rarely in heads, sub-

tended by one or more bracts, which often enclose the

young inﬂ orescence; individual ﬂ owers sessile and

often even embedded in the axis or rarely shortly

pedicellate, bisexual or much more often unisexual

(monoecious or less often dioecious) or sometimes

polygamous, actinomorphic or less often weakly

zygomorphic, commonly 3-merous, usually bracteate.

Perianth segments six or sometimes four, in two more

or less dissimilar cycles or less often up to ten and spi-

rally arranged, free or more or less connate, often scale-

like, rarely reduced or lacking; outer segments (sepals)

usually smaller, imbricate or open in bud; inner seg-

ments (petals) valvate in male ﬂ owers and imbricate in

female ﬂ owers. Stamens mostly six in two cycles, but

seldom, as in Nypa, one of two cycles lacking, or some-

times numbers range from seven to eight to mostly nine

to many (120 to over 250 in Phytelephas, up to 521 in

Ammandra and up to 954 in Palandra), ﬁ laments free

Subclass III. ARECIDAE

694 Subclass III. ARECIDAE

or more or less connate or/and adnate to the inner peri-

anth segments; anthers tetrasporangiate, basiﬁ xed or

dorsiﬁ xed, latrorse, opening usually longitudinally or

rarely (some species of Areca) by apical pores. Tapetum

secretory. Microsporogenesis simultaneous or some-

times successive (reported for Nypa fruticans, Phoenix

sylvestris, and Pinanga disticha, but needs conﬁ rmation).

Pollen grains tectate-columellate, 1(2)-colpate, tricho-

tomocolpate, zonocolpate or 1(2)-porate, predominantly

reticulate. Staminodia often present. Gynoecium of

(1)3(4–10) free or more often united carpels, some-

times pseudomonomerous; stylodia free or connate;

ovary superior, in syncarpous forms usually 3-locular,

with one erect or pendulous ovule per locule or per

carpel. Septal nectaries often present. Ovules anatro-

pous or less often hemitropous, campylotropous, or

orthotropous, bitegmic, crassinucellate, with parietal

cell. Female gametophyte usually of Polygonum-type,

rarely of Allium-type. Endosperm nuclear. Fruits dry or

ﬂ eshy drupe with the endocarp usually attached to the

seed or less often baccate, rarely more or less dehis-

cent. Seeds with very small, cylindric, or conical

embryo embedded apically, laterally or basally in copi-

ous endosperm; endosperm usually bony, sometimes

ruminate, contains fats and oils, aleurone and hemicel-

lulose deposited in cell walls, but without starch.

n = 8–10, 12–19, especially 14, 16, 18. In Voanioala

2n = 606 ±3 (Johnson 1989; Roser 1994).

Arecales are one of the most ancient branches of the

monocotyledons and are among the ﬁ rst families of

magnoliophytes that are deﬁ nitely recognizable in the

fossil record. They are also very heterobathmic. There

are 16 arecaceous genera with the apocarpous gynoe-

cium, and, what is even more important, in some of

them the carpels are conduplicate and stipitate, with

open sutures and laminar or sublaminar placentation

(see Uhl and Moore 1971; Moore 1973). In

Trachycarpus fortunei trichomes are present along and

in some extent within the unsealed suture, as in some

taxa of archaic dicotyledons (Uhl and Moore 1971;

Moore and Uhl 1982). Stigmas are sessile or sub-

sessile. In Arecales septal nectaries developed inde-

pendently, and as Eames (1953) says “the story here

parallels that in the Liliaceae; septal glands have devel-

oped independently, perhaps several times.”

The Arecales resemble the commelinalean and

poalean families in possessing epicuticular wax of the

Strelitzia-type and cell walls containing UV-ﬂ uorescent

compounds. Besides, the Arecales resemble the

Joinvilleaceae, Flagellariaceae, and Poaceae in plicate

ptyxis of leaves (found in Joinvillea and some grasses),

silica bodies, the occurrence of vessels in the stems,

the pronounced development of ligules and ligulelike

structures, the conspicuously similar ﬂ avonoid com-

pounds, such as tricin and sulfonated ﬂ avonoids, and

abundant cyanogenesis (Dahlgren and Rasmussen

1983; Dahlgren et al. 1985). But in contrast to the

Poales, septal nectaries are common in palms, and,

what is phylogenetically more important, some of

them still have such plesiomorphic characters as

apocarpous gynoecium of very primitive carpels and

simultaneous microsporogenesis. In general, palms are

much nearer to the archaic Liliidae and Alismatidae

than to the Poanae and related groups. However, they

occupy a very isolated taxonomic position, and evi-

dently originated independently and directly from the

earliest monocotyledons.

1. ARECACEAE

Schultz-Schultzenstein 1832 or Palmae A. L. de

Jussieu 1789 (nom. altern.) (including Acristaceae

O. F. Cook 1913; Borassaceae Schultz-Schultzenstein

1832; Calamaceae Kunth ex Perleb 1838; Caryotaceae

O. F. Cook 1913; Ceroxylaceae O. F. Cook 1913;

Chamaedoraceae O. F. Cook 1913; Cocosaceae

Schultz-Schultzenstein 1832; Coryphaceae Schultz-

Schultzenstein 1832; Geonomataceae O. F. Cook 1913;

Iriarteaceae O. F. Cook et Doyle 1913; Lepidocaryaceae

C. Martius 1838; Malortieaceae O. F. Cook 1913;

Manicariaceae O. F. Cook 1910; Nypaceae Brongniart

ex Le Maout et Decaisne 1868; Phoenicaceae Burnett

1835; Phytelephantaceae Martius ex Perleb 1838;

Pseudophoenicaceae O. F. Cook 1913; Sabalaceae

Schultz-Schultzenstein 1832; Sagaceae Schultz-

Schultzenstein 1832; Saginaceae Berchtold et J. Presl

1820; Synechanthaceae O. F. Cook 1913). 188/2000–

2700. Pantropical, with some subtropical and a few

temperate species.

Classiﬁ cation after J. Dransﬁ eld et al. 2005 and

M.V. Norup et al. 2006.

1.1 CALAMOIDEAE

Erect or climbing, often spiny. Leaves palmate,

costapalmate, or, more often, pinnate, reduplicate.

Flowers almost always in dyads or dyad derivatives,

bisexual, polygamous, monoecious or dioecious, with