Takhtajan Armen. Flowering Plants

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Superorder LILIANAE 635

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Order 9. BURMANNIALES

Mostly small, perennial or annual mycorrhizal herbs,

achlorophyllous and mycotrophic or less often auto-

trophic. The perennial autotrophic members with elon-

gate or sometimes tuberous rhizome or tubers. Vessels

with scalariform perforations present in roots, stems,

and leaves of autotrophic species of Burmannia. Sieve-

element plastids of P2c-type (Behnke 2002). Leaves

alternate, basally sheathing, in mycotrophic species

more or less reduced to scales, in autotrophic species

linear to lanceolate and mostly concentrated at the base

of the stem. Stomata anomocytic. Flowers terminal, in

cymose inﬂ orescences (usually a bifurcate cincinnus)

or solitary, bisexual, actinomorphic to more or less

zygomorphic, 3-merous. Perianth of 6(-8) segments in

two cycles, more or less connate into a tube, sometimes

the inner cycle absent. Stamens six in two cycles or

(Burmanniaceae and rarely in Thismiaceae) only three

and then opposite the inner segments (in contrast to the

Iridales, where the stamens alternate with the inner

segments); ﬁ laments usually short, attached to the peri-

anth tube or (Corsiaceae) free from the perianth; anthers

mostly with short ﬁ laments, tetrasporangiate, opening

longitudinally or laterally, extrorse or introrse,. Tapetum

secretory. Microsporogenesis successive. Pollen grains

mostly 3-celled, tectate-columellate, 1-colpate or 1–2-

porate, sometimes polyforaminate or inaperturate, psi-

late, undulate, folded or ﬁ nely reticulate. Gynoecium

of three united carpels, with ﬁ liform or short, simple or

apically 3-brachiate style; ovary inferior, 3-locular to

1-locular, with numerous ovules per locule or per pari-

etal placenta. Septal nectaries present or absent. Ovules

minute, anatropous, bitegmic, tenuinucellate, without

parietal cell. Female gametophyte of Polygonum-type

or sometimes (in some species of Burmannia) of

Allium-type. Endosperm helobial or rarely (some

species of Thismia) cellular. Fruits usually capsular

and dehisce longitudinally or transversely by slits or by

valves, with terminal aperture or irregularly, rarely

indehiscent. Seeds numerous and very small, with min-

ute and undifferentiated, mostly globular embryo that

often consists of 3–8 cells; endosperm usually consists

of few cells; testa usually reticulate.

According to Dahlgren et all (1985: 216) “The clos-

est relatives are perhaps the Melanthiales, in which

Campynemataceae takes an intermediate position

between the orders”. Rudall and Eastman (2002) also

came to the conclusion, that there are range of mor-

Superorder LILIANAE 637

phological similarities between Corsia and Campy-

nemataceae, which in their opinion tends to support

preliminary inferences from molecular data that they

are closely related. Thus, Burmanniales are most prob-

ably closely related to Campynemataceae, which in

my system is placed in Liliales.

Key to Families

1 Fertile stamens three. Autotrophic or saprophytic,

annual or perennial herbs. Vessels, when present, in

roots and stems, with scalariform perforations.

Leaves well developed, or much reduced, or absent,

basal or with neither basal nor terminal aggrega-

tions of leave, alternate, spiral or distichous, sessile,

simple, entire, linear to ovate. Plants without silica

bodies. Flowers solitary or in terminal cymes.

Perianth generally actinomorphic, with straight or

rarely curved tube and often very short lobes; often

both perianth tube and ovary 3-angled or 3-winged.

Stamens opposite the inner segments (when pres-

ent); ﬁ laments short, inserted in the perianth tube;

anthers introrse to latrorse, with thick and broad,

sometimes transversely extended connective, open-

ing transversely. Pollen monoporate or inaperturate,

psilate. Nectaries septal at the top of ovary or dis-

persed on its side. Style apically divided into three

branches, each of which bears stigmatic surface;

ovary 3-locular and with protruded axile placentas

or 1-locular with parietal placentas, which may

emerge centrally at the ovary base. Ovary anatro-

pous, tenuinucellate. Fruits capsules, loculicidal or

rarely valvular. Seeds with testa extended into long

wing at each end; embryo rudimentary at the time

of seed release, testa without phytomelan; n = 6–8,

much and high polyploidy. . . .1. burmanniaceae.

1 Fertile stamens usually six. Plants always

mycotrophic. Nectaries absent.

2 Stamens inserted in perianth tube, usually six, but

in Oxygyne three, recurved to pendent; thecae lon-

gitudinally dehiscent and united (separate when

forming an anther tube), introrse. Small, often

somewhat succulent, perennial, rhizomatous

herbs; rhizome sometimes branched and coral-

like. Leaves alternate or opposite, small, distic-

hous, sessile, simple, entire. Mesophyll containing

calcium oxalate crystals. Flowers in a few-ﬂ ow-

ered cincinnus or more often solitary, small, zygo-

morphic or actinomorphic. Perianth segments

similar in the two whorls, or different (the lobes

equal, or those of the inner whorl longer). Anthers

basiﬁ xed, appendages (with apical enlargement of

the connective. Tapetal cells unicellular, pollen

porate. Ovary 1-locular, with three parietal pla-

centas; styles one, apical, stigmas three, placenta-

tion parietal. Ovules numerous, anatropous. Fruits

circumscissile capsules. Seeds numerous, minute,

embryo rudimentary at the time of seed release,

tecta without phytomelan; endosperm lacking; n =

6–8, 11–13.. . . . . . . . . . . . . . . . . 2. thismiaceae.

2 Stamens free from the perianth. Mycotrophic

herbs, erect or (Corsia) creeping with rhizomes

or tubers. Plants without silica bodies. Leaves

alternate, sessile, simple, entire, reduced to rather

large scales, spiral to 2-ranked. Flowers solitary,

terminal, bisexual or (Corsiopsis) dioecious or

monoecious. Perianth strongly zygomorphic, one

of the segments (the posterior of the outer cycle)

large and brightly coloured (red-violet), adaxial;

the other linear- ﬁ liform and more or less reﬂ exed.

Stamens six; ﬁ laments short; anthers dorsiﬁ xed,

extrorse. Pollen grains sulcate or ulcerate. Ovary

inferior, 1-locular, with three much intruded and

bifurcate placentas. Stylodia free or connate into

a style; stigmas thick. Ovules numerous, anatro-

pous, tenuinucellate. Endosperm helobial. Fruits

capsules dehiscing by three vertical valves down

to the base or with terminal aperture. Seeds

numerous, minute or small, with reticulate testa

prolonged into a wing at each end and testa with-

out phytomelan; embryo rudimentary at the time

of seed release; endosperm multicellular (Corsia)

or reduced to few cells (Arachnitis), oily or not

oily; n = 9.. . . . . . . . . . . . . . . . . .3. corsiaceae.

1. BURMANNIACEAE

Blume 1827. 10/95. Widely distributed in the tropics

and subtropics.

Burmannia, Campylosiphon, Hexapterella, Dicty-

ostega, Miersiella, Cymbocarpa, Gymnosiphon,

Apteria, Marthella, Geomitra.

2. THISMIACEAE

J. Agardh 1858. 5/40. Southeastern Asia, Africa,

America (mainly Brazil), Australia, and New Zealand.

Oxygyne, Afrothismia, Haplothismia, Thismia,

Scaphiophora.

638 Subclass II. LILIIDAE

3. CORSIACEAE

Beccari 1878 (including Arachnitidaccae Munoz

1966). 3/30. China (Corsiopsis), New Guinea, Solomon

Islands, and Australia (Corsia, 27) and Bolivia, Chile,

Patagonia and Falkland Islands (Arachnitis, 2).

Corsia, Corsiopsis, Arachnitis.

Close related to the Burmanniaceae.

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Order 10. ALSTROEMERIALES

Erect or sometimes climbing rhizomatous perennial

herbs or vines with ﬁ brous and often also fusiform and

ﬂ eshy tuberlike roots containing starch. Stems leafy,

those of climbing species up to 4–6 m long. Calcium

oxalate raphides present. Vessels both in roots (mostly

with scalariform perforations) and stems (only with

scalariform perforations). Axial parenchyma contains

starch. Leaves alternate, linear to oblong, glabrous or

pubescent on both leaf surfaces; epicuticular wax

sometimes present. Stomata anomocytic. Inﬂ orescences

terminal or axillary, consisting of helicoid cymes,

umbel-like or capitate, rarely ﬂ owers solitary. Flowers

usually few to numerous (up to 100 in some Bomarea),

bisexual, more or less actinomorphic or zygomorphic

Superorder LILIANAE 639

(Alstroemeria), 3-merous. Perianth segments six, in two

cycles, usually free. Stamens six in two cycles; ﬁ la-

ments free, narrow; anthers basiﬁ xed, elongate, introrse

or extrorse, tetrasporangiate, opening longitudinally,

rarely by apical pores (Luzuriagaceae). Tapetum

secretory. Microsporogenesis successive. Pollen grains

2-celled, large, 1-colpate, usually plano-convex.

Gynoecium of three united carpels; style ﬁ liform, more

or less 3-lobed; ovary inferior or superior (Lazuriagaceae),

3-locular or 1-locular. Ovules anatropous, bitegmic,

tenuinucellate or (Petermannia) crassinucelate Female

gametophyte of Polygonum-type. Endosperm nuclear.

Fruits loculicidal capsules, or berries. Seeds globose to

ellipsoid, embryo straight, cylindrical or linear; seeds

lacking phytomelan; endosperm copious, contains

hemicellulose, aleurone, and fatty oils; Steroidal

saponins present or absent; contain chelidonic acid,

ﬂ avonol glycosides; n = 5, 8, 9, 10.

Most probably related to the Liliaceae, which is

also supported by the accumulation of appreciable

amounts of tuliposide A (Slob et al. 1975). Hallier

(1912) derived (with a question mark) Alstroemeriaceae

from the Liliaceae.

Key to Families

1 Ovary superior.

2 Steroidal saponins present. Slender shrubby

perennials from short rhizome. Calcium oxalate

raphides present. Roots ﬁ brous. Rhizome with

monocotyledonous secondary thickening. Stems

thin, branching, erect or twisting. Vessels with

simple or scalariform perforations. Leaves alter-

nate, distichous, shortly petiolate, ovate, parallel-

veined, with many cross-veins. Stomata

anomocytic. Flowers in axillary reduced cymes

(bostryx), or solitary, functionally dioecious,

small, hypogynous, actinomorphic, articulated

on the pedicels with a short pericladium. Perianth

segments six, not twisted after anthesis, united

about two thirds of their length into a campanu-

late tube, persistent. Stamens six in two whorls;

ﬁ laments adnate to the tepals; anthers 2-locular,

tetrasporangiate, basal- dorsiﬁ xed, introrse. In

female ﬂ owers stamens adnate to the perianth

segments and reduced in size. Pollen grains obo-

vate and sulcate, with reticulate exine. Gynoecium

of three united carpels; style with 3-lobed, wet

stigma; ovary 3-locular; ovules 2–3 per locule,

anatropous, tenuinucellate; in male ﬂ owers ovary

is shorter, lacks loculi, but has well- developed

septal nectaries. Fruits berries; seeds angular,

without phytomelan; testa and tegmen thin-

walled; endosperm copious, storing aleurone and

lipids; embryo clavate, about two thirds as long

as the seed. . . . . . . . . . . . . . . . . 2. behniaceae.

2 Steroidal saponins absent. Terrestrial or epiphytic

perennials with thin woody stems and thin rhi-

zomes (Drymophila) or shrubs with short rhi-

zomes (Luzuriaga). Stems thin, branching or

simple, erect or scrambling. Vessels in the roots

and stems, with simple or scalariform perfora-

tions. Calcium oxalate raphides and druses pres-

ent in Drymophila. Leaves alternate, distichous,

sessile or slightly petiolate, simple, linear to ovate,

with numerous parallel veins and very slight retic-

ulate or transverse venation, lamina inverted; mes-

ophyll containing mucilage cells and calcium

oxalate crystals. Flowers bisexual, in axillary,

monobrachial cincinnus or solitary, actinomor-

phic, articulated on the pedicels (Luzuriaga) or at

base of ovary (Drymophila). Perianth segments

six or rarely eight in Drymophila, free almost to

base, nectariferous at the base. Stamens six in two

cycles; ﬁ laments free; anthers basiﬁ xed or dorsi-

ﬁ xed, introrse or extrorse, opening longitudinally

or sometimes by apical pores. Pollen grains 1-col-

pate, reticulate, or (Geitonoplesium) trichotomo-

sulcate. Gynoecium of three united carpels; style

with deeply 3-lobed (Drymophila) or capitate

stigma; ovary superior 3-locular or rarely 1-locu-

lar, with several anatropous to hemitropous ovules.

Fruits berries or berry-like, ﬂ eshy capsules. Seeds

few to numerous, small, testal, in Eustrephus with

ﬂ eshy strophiole. Endosperm copious, storing fat

and aleurone and also hemicellulose (except

Geitonoplesium); embryo straight, usually rather

long, or curved (Geitonoplesium, Eustrephus).

n = 10. . . . . . . . . . . . . . . . . . 1. luzuriagaceae.

1 Ovary inferior.

3 Fruits capsules or rarely (some sp. Bomarea) fruits

berrylike and indehiscent. Erect or sometimes

climbing rhizomatous perennial herbs or vines

(most Bomarea), with ﬁ brous and often also fusi-

form and ﬂ eshy tuberlike roots containing starch;

very rarely (Alstroemeria graminea) annuals with-

out storage roots. Stems unbranched, leafy, those

of climbing species up to 4 m long or even longer.

Calcium oxalate raphides present. Vessels both in

640 Subclass II. LILIIDAE

roots (mostly with scalariform perforations) and

stems (only with scalariform perforations). Axial

parenchyma contains starch. Leaves alternate, lin-

ear to oblong, narrowing at the base, generally

twisted at the base by 180 degrees and reversing

the surfaces, glabrous or pubescent on both leaf

surfaces; epicuticular wax sometimes present.

Stomata anomocytic. Inﬂ orescences terminal,

consisting of helicoid cymes, umbel-like or

(Leontochir) capitate, rarely uniﬂ oral. Flowers

usually few to numerous (up to 100 in some

Bomarea), rather large, subtended by relatively

large leaf-like bracts, bisexual, more or less acti-

nomorphic or zygomorphic (Alstroemeria), 3-mer-

ous. Perianth segments six, in two cycles, usually

free, more or less similar to conspicuously differ-

ent in two cycles (the inner often longer, of differ-

ent colour and more variegated than the outer

one), in all genera the base of all or two of the

inner tepals is canaliculate and in Leontochir even

closed to form a pocket and contains nectar from

succulent trichomes (Bayer 1998). Stamens six in

two cycles; ﬁ laments free, narrow; anthers pseudo-

basiﬁ xed, elongate, sagittate, tetrasporangiate,

opening longitudinally, latrorse. Tapetum secre-

tory. Pollen grains 2-celled, large, 1- colpate, usu-

ally plano-convex, semitectate. Gynoecium of

three united carpels; style ﬁ liform, more or less

3-lobed; ovary inferior, 3-locular or 1-locular

(Leontochir and one species of Alstroemeria by

abortation late in development), with numerous

ovules per locule or per parietal placenta; distinct

decurrent borders of the outer perianth segments

are seen on the surface of the ovary as well as a

circular scar around the top of the fruit left after

the shedding of the perianth. Placentation axile

(Alstroemeria, Bomarea) or parietal (Leontochir,

Alstroemeria pygmaea). Ovules anatropous, tenu-

inucellate, with funicular obturator; no parietal

cell cut off from the archesporial cell. Fruits more

or less truncate capsules, opening loculicidally

from the top to the base or only on the top, in

Alstroemeria mostly explosively, rarely (some

species of Bomarea) fruits berrylike and indehis-

cent. Seeds globose or globose to ellipsoid, mostly

tuberculate, with long, narrow raphe and well-

expressed hilum; testa of few to many cell layers,

dry or (Bomarea) ﬂ eshy, with small amount of

phlobaphenes in endotesta (at least in Alstroemeria),

the inner integument reduced to thin membrane;

endosperm copious, consists of cells with large

pores, contains hemicellulose, aleurone, and fatty

oils; embryo straight, cylindrical, about one-third

to two-thirds the length of the endosperm. Produce

steroidal saponins and chelidonic acid as well as

ﬂ avonol glycosides and tuliposides (in which the

order agrees with the Liliaceae), but alkaloids are

lacking, n = 8 (Alstroemeria) or 9 (Leontochir and

Bomarea). Chromosomes generally large. . . . . . .

3. alstroemeriaceae.

3 Fruits berries. Perennial, rhizomatous, woody

lianas with ﬁ brous irregularly distributed

adventitious roots and more or less prickly,

climbing or scrambling stems to 6 m long.

Calcium oxalate raphides present in all parts.

Branching sympodial and some of morphologi-

cally terminal inﬂ orescences transformed into

branched tendrils. Vessels in roots and stem,

with scalariform perforations. Leaves alternate,

shortly petiolate, broadly lanceolate, acumi-

nate, pinnate- striate veined with reticulate ﬁ ner

venation. Flowers in leaf-opposed or terminal

branched cymes, bisexual, not articulated,

ebracteate. Perianth segments six, subequal,

spreading, or reﬂ exed, basally nectariferous.

Stamens six, free, inserted at the base of peri-

anth; ﬁ laments ﬁ liform; anthers oblong, bas-

iﬁ xed, tetrasporangiate, extrorse. Tapetum

plasmodial. Pollen grains 1-colpate, with small

spinules, contains starch grains at anthesis

(Conran and Clifford 1986). Gynoecium of

three carpels; style slender, with capitate,

slightly 3-lobed stigma; ovary 1-locular, with

numerous anatropous, crassinucellate ovules

on parietal placentas. Berries bright red, with

thick, ﬂ eshy pericarp. Seeds numerous more or

less globose but somewhat angular, rugulose,

dark brown, with thickened exotesta, several-

layered mesotesta and thick-walled endotesta;

tegmen consists of collapsed thin-walled cells;

embryo linear, medium-sized; endosperm copi-

ous. Steroidal saponins and alkaloids lacking,

n = 5. . . . . . . . . . . . . . . 4. petermanniaceae.

1. LUZURIAGACEAE

J.P. Lotsy 1911 (including Eustrephaceae Chupov

1994, including Geitonoplesiaceae R. Dahlgren ex

Conran 1994). 4/7. South America from Peru to Tierra

Superorder LILIANAE 641

del Fuego, and Falkland Islands; Southern Africa,

Malesia from Java to New Guinea, Australia

(New South Wales to Tasmania), New Zealand,

New Caledonia.

luzuriageae: Drymophila, Luzuriaga;

eustrepheae: Eustrephus, Geitonoplesium

2. BEHNIACEAE

Conran, Rudall et Chase 1997. 1/1. Southern and

South-Eastern Africa.

Behnia.

Taxonomic position of monotypic genus Behnia is

debatable. Schlittler (1953) included this genus in

Luzuriagoideae (Liliaceae s.l.). The molecular studies

of Chase et al. (1995) found instead that Behnia occu-

pied a basal position to clade containing the Agavaceae,

Anthericaceae.

3. ALSTROEMERIACEAE

Dumortier 1829. 3/280. From central Mexico and West

Indies to Chile and Argentina, mostly in the Andes.

Leontochir, Bomarea, Alstroemeria (including

Schickendantzia and Taltalia).

Related to the Luzuriagaceae (Bayer 1998; Meerow

2004), which is also supported by the presence of

colchicines alkaloids, ﬂ avonol glycosides (and absence

of ﬂ avones) and the accumulation of appreciable

amounts of tuliposide A (Hegnauer 1963; Slob et al.

1975), as well as by the molecular and morphological

data (Chase et al. 1995; Rudall et al. 2000; Vinnersten

and Bremer 2001; Soltis et al. 2006).

4. PETERMANNIACEAE

Hutchinson 1934. 1/1. Eastern Australia.

Petermannia.

Closely related to the Luzuriagaceae (Stevenson

and Loconte (1995)

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Order 11. SMILACALES

Rhizomatous herbs or shrubs. Rhizomes with or

without secondary thickening. Roots ﬁ brous. Oxalate

raphides present. Vessels in roots, in roots and stem, or

sometimes also in leaves, with scalariform, simple, or

scalariform and simple perforations. Leaves alternate,

distichous, opposite or verticillate, sessile or petiolate,

mostly without a sheathing base, mostly coriaceous,

with several prominent longitudinal veins, often with

usually prominent reticulate or transverse venation;

midrib or main veins prominent. Stomata anomocytic.

Flowers in terminal or axillary inﬂ orescences or rarely

solitary, mostly articulated on the pedicels, bisexual

(Philesiaceae, and Ripogonaceae) or unisexual and

dioecious, actinomorphic, 3-merous. Perianth seg-

ments six, usually more or less similar, free or connate

into a tube. Stamens six, rarely (some Smilacaceae)

three or nine (up to 18); ﬁ laments usually free or some-

times (Heterosmilax) more or less connate into a tube

or column, sometimes adnate to the base of perianth

segments or to the perianth tube; anthers basiﬁ xed or

dorsiﬁ xed, tetrasporangiate or disporangiate, extrorse,

latrorse or introrse, usually opening longitudinally.

Tapetum secretory or (Ripogonum) plasmodial.

Microsporogenesis successive. Pollen grains 2-celled,

1-colpate and with reticulate ornamentation or inaper-

turate and spinulose. Nectaries often occur at the bases

of perianth segments or stamens. Gynoecium of three

united carpels; stylodia free or only basally connate or

connate into a style with 3-lobed or capitate stigma;

ovary superior 3-locular or 1-locular, with one or two

to numerous ovules per locule or per parietal placenta.

Septal nectaries absent. Ovules anatropous, hemitrop-

ous, campylotropous, or orthotropous, bitegmic,

crassinucellate, with parietal cell. Female gametophyte

of Polygonum-type. Endosperm nuclear. Fruits berries.

Seeds without phytomelan crust, with straight and

sometimes very small embryo and copious endosperm

containing aleurone, lipids, and sometimes hemicellu-

lose, rarely (Ripogonum) starch. Lacking alkaloids.

Related to the Alstroemeriales and probably derived

from the same stock.

Key to Families

1 Plants without tendrils.

2 Leaves alternate, usually with inverted lamina.

Erect (Philesia) or climbing (Lapageria) shrubs

Superorder LILIANAE 643

from short woody rhizomes. Vessels in roots and

stem, with scalariform perforations. Leaves ses-

sile or petiolate, more or less sheathing at base,

ovate to lanceolate or broadly linear and twisted

at base, but lamina not inverted, with one

(Philesia) or 3–5 (Lapageria) prominent parallel

veins and distinctly reticulate or transverse vena-

tion. Stomata anomocytic (in Lapageria trans-

versely oriented in relation to parallel veins).

Flowers solitary in leaf axils or terminal on short

branchlets; pedicels not articulated. Perianth seg-

ments free, in Philesia the outer cycle short and

nearly sepaloid; inner segments always large,

petaloid and overlapping to form a campanulate

perigone. Filaments free (Lapageria) or connate

half-way up (Philesia), anthers subbasiﬁ xed,

extrorse or introrse, opening longitudinally.

Pollen grains inaperturate and spinulose. Style

linear, with 3-lobed or capitate stigma; ovary

1-locular, with numerous anatropous or amphi-

tropous ovules on intrusive parietal placentas.

Seeds globose; endosperm copious and consists

of cells with rather thin, nonpitted walls; embryo

1/2 as long as the seed. Steroidal saponins pres-

ent. n = 15, 19. . . . . . . . . . . . . 1. philesiaceae.

2 Leaves opposite, decussate or verticillate. Lamina

noninverted. Commonly climbing shrubs with

glabrous or pubescent and often prickly stems,

sometimes up to 20 m tall. Vessels only in roots

and stem, with scalariform perforation. Leaves

with twisted petiole, 3–5 parallel veins and retic-

ulate transverse venation. Flowers in axillary

racemes or spikes or in terminal racemose pani-

cles, usually bisexual. Perianth segments free,

similar, spreading. Filaments free, very short;

anthers basiﬁ xed or subbasiﬁ xed, tetrasporan-

giate, opening longitudinally, latrorse. Pollen

grains 1- colpate, reticulate and slightly foveolate.

Style very short and thick or absent; stigma

minutely 3-lobed; ovary 3-locular, with (1)2 ana-

tropous ovules per locule. Seeds brown; seed coat

derived mainly from tegmen and chalazal layers,

whereas ﬂ eshy testa are represented by cuticular

membrane that includes obliterated remnants of

outer integument; endosperm copious, very hard,

consists of radially elongated, thick-walled cells

with scalariform-pitted walls, contains starchy

grains and hemicellulose; embryo very small,

linear. n = 15. . . . . . . . . . . . . .2. ripogonaceae.

1 Plants mostly with paired tendrils on petiole. Woody

or herbaceous creeping, climbing, or scrambling

plants with rhizomatous or tuberous rootstock and

often prickly stems and branches. Vessels in all

parts, with scalariform perforations. Leaves alter-

nate, lower ones reduced to scales, upper ones lan-

ceolate to elliptic or cordate-orbiculate, petiolate,

entire or slightly lobed, 3–9-veined, reticulate-veiny

between parallel veins. Flowers rather small, in axil-

lary, usually peduncled umbels or fascicles that are

solitary or arranged on elongated axis, dioecious.

Perianth segments free (Smilax) or united into den-

tate tube. Stamens six (rarely up to 15) or less often

nine (Pseudosmilax) or only three (Heterosmilax)

free or (Heterosmilax) united into a column; anthers

basiﬁ xed, disporangiate, introrse. Pollen grains in

monads, globose, inaperturate or indistinctly 1-col-

pate, spinulose. Male ﬂ owers without pistillodia.

Stylodia short, thick, and spreading, free or rarely

basally shortly connate, papillate; ovary 3-locular,

with 1–2 pendulous, orthotropous ovules. Staminodia

in female ﬂ owers present. Fruits berries. Seeds

brown, shiny; when ripe, testa is obliterated and only

tegmen remains, phlobaphenes; endosperm copious,

consists of radially elongated, thick-walled cells and

contains hemicellulose, aleurone, and fatty oils, but

starch is lacking; embryo small, linear, straight. n =

13, 14, 15, 16. . . . . . . . . . . . . . . . 3. smilacaceae.

1. PHILESIACEAE

Dumortier 1829. 2/2. Southern Chile.

Philesia, Lapageria.

Closely related to the Smilacaceae (see Conran

1989). Molecular studies also supported a close rela-

tionship between the Philesiaceae and the Smilacaceae

(Chase et al. 1995; Soltis et al. 2006).

2. RIPOGONACEAE

Conran et H. Clifford 1985. 1/6. New Guinea, eastern

Australia, New Zealand, and Stewart and Chatham

Islands.

Ripogonum.

Related to the Philesiaceae and in some respects

occupying an intermediate position between them and

the Smilacaceae.

644 Subclass II. LILIIDAE

3. SMILACACEAE

Ventenat 1799. 2/320–370. Mainly tropical and sub-

tropical regions, but also in warm-temperate regions.

Smilax, Heterosmilax (Pseudosmilax).

Close to the Ripogonaceae and the Philesiaceae.

All this families have unique spiny pollen (Rudall

et al. 2000).

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Order 12. ORCHIDALES

Perennial herbs, terrestrial or epiphytic, usually

mycotrophic or occasionally saprophytic, rhizomatous

or tuberous. Silicia bodies enclosed in special cells,

calcium oxalate raphides usually present. Vessels

mostly only in roots, sometimes also in the stem and

also in leaves, mostly with scalariform perforations.

Leaves alternate or rarely opposite, parallel-veined.

Stomata mostly paracytic or sometimes tetracytic.

Flowers in various types of inﬂ orescences, usually

bisexual, mostly zygomorphic. Perianth segments six,

free or more or less connate. Stamens six, three, two or

usually one. Anthers dorsiﬁ xed or basiﬁ xed, introrse

or seldom latrorse, tetrasporangiate, opening longitu-

dinally. Tapetum secretory or rarely plasmodial.

Microsporogenesis simultaneous or successive. Pollen

grains mostly 1-colpate, tectate-columellate.

Gynoecium of three united carpels; ovary superior or

more often inferior, 3-locular or rarely unilocular,

ovules of various number but mostly numerous, anat-

ropous, very rarely hemitropous or campylotropous,

crassinucellate or more often tenuinucellate, usually

bitegmic. Female gametophyte of Polygonum-type or

rarely of Allium-type, very rarely tetrasporic.

Endosperm mostly nuclear. Fruits mostly capsular.

Seeds usually numerous. Endosperm reduced or more

often lacking, very rarely (Lanaria) copious. Embryo

well developed or more often reduced and

undifferentiated.

Very old group, which probably originated from the

melanthianean ancestor.