Takhtajan Armen. Flowering Plants

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A. Takhtajan, Flowering Plants, 625

Perennial or annual herbs or more or less arborescent

plants often with secondary growth of monocotyledon-

ous type, often with well-developed subterranean

storage organs, such as rhizomes, bulbs, corms, and

tubers. Vessels present or absent, with scalariform or

simple perforations. Leaves both basal and cauline or

only basal, alternate, opposite, or verticillate, mostly

with parallel venation. Stomata of various types.

Flowers from very small to rather large in various

kinds of inﬂ orescences or solitary, bisexual or unisex-

ual, actinomorphic or zygomorphic, with double or

simple perianth, entomophilous or anemophilous.

Tapetum usually secretory. Microsporogenesis mostly

successive. Pollen grains 2-celled or 3-celled, 1- colpate

or of various derived types. Gynoecium apocarpous or

syncarpous. Ovules mostly anatropous, commonly

bitegmic, usually crassinucellate. Female gametophyte

mostly of Polygonum-type. Endosperm helobial or

nuclear. Seeds mostly with more or less copious

endosperm (lacking in Orchidales).

The most archaic liliids are characterized by some

very primitive features, especially in their carpel and

pollen morphology. However, some members of the

Liliidae, including Orchidaceae, have reached a very

high level of specialization.

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Subclass II. LILIIDAE

626 Subclass II. LILIIDAE

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Superorder LILIANAE

Order 5. MELANTHIALES

Perennial or rarely annual herbs, commonly with rhi-

zomes, rarely cormlike, and sometimes with a bulblike

base but without nutrient bulb scales. Raphides often

present. Roots ﬁ brous. Vessels only in roots, with sca-

lariform perforations. Aerial stem erect, simple, stout

to slender, with foliage or scaly leaves, sometimes

basally thickened. Leaves deciduous or evergreen,

alternate, all cauline (Veratrum spp.), or with large

basal and small cauline leaves (Xerophyllum, Zigadenus

spp.), or all leaves in a basal rosette (Heloniopsis),

bifacial, distichous, sheathing at the base, linear, lan-

ceolate, or rarely ovate, up to 50 cm long. Stomata ano-

mocytic. Inﬂ orescences terminal, often racemes,

sometimes panicles, spikes or umbel-like, glabrous or

pubescent. Flowers small, bracteate or ebracteate,

without bracteoles, bisexual or very rarely unisexual,

3-merous, actinomorphic or rarely (Chionographis)

zygomorphic. Perianth segments six in two cycles,

generally similar, free or sometimes more or less

basally connate, in some genera with basal nectaries.

Stamens usually six in two cycles; ﬁ laments free or

rarely adnate to the base of the perianth segments; ﬁ la-

ments ﬁ liform to subulate; anthers tetrasporangiate,

basiﬁ xed or dorsiﬁ xed-hypopeltate, mostly extrorse,

opening longitudinally, Tapetum secretory.

Microsporogenesis successive. Pollen grains 2-celled,

tectate-columellate or less often (Heloniadeae and

Chionographideae) intectate, mostly 1-colpate, or

4-porate (Chionographideae). Gynoecium of three

carpels, free or more or less united and with free or

connate stylodia; ovary superior or less often semi-

inferior; each carpel, each locule and each parietal pla-

centa with two to numerous ovules. Ovules anatropous

or sometimes campylotropous and epitropic, bitegmic,

crassinucellate to tenuicellate, with parietal cell.

Female gametophyte of Polygonum-type or

Allium-type. Endosperm helobial. Fruits septicidal

Chionographideae), loculicidal (Xerophylleae and

Heloniadeae), or ventricidal capsules, conical to

globular, often 3-lobed. Seeds generally provided with

terminal appendages or wings; seed coat essentially

testal; the testal part has few cell layers, lack phytomelan

but sometimes has phlobaphene; the tegminal part is

thin and collapsed, red brown, or yellowish, though

ﬂ attened; endosperm usually consists of thin-walled

cells containing aleurone and fatty oils, sometimes

also rounded starch grains; embryo small (1/5–1/9 of

the length of endosperm), basal, globose or ovoid, or

rather large and almost equal to the length of the seed

(as in species of Melanthium and Zigadenus) and axial

and linear. Steroidal saponins and chelidonic acid

occur in many genera. Veratrum and some other related

genera contain alkaloids derived from steroidal precur-

sors; n = 6 (Chionographis – Zomlefer 1997)

n = 8, 10–12, 15–17, 21–22.

Melanthiales, especially genera with more or less

free carpels, are the most archaic members of the

Liliidae and probably of the monocotyledons in general

(see Lotsy 1911; Takhtajan 1959, 1966, 1980, 1987;

Eames 1961; Radulescu 1973d).

1. MELANTHIACEAE

Batsch 1802 (including Chionographidaceae Takhtajan

1996; Heloniadaceae J. Agardh 1858; Veratraceae Salis-

bury 1807; Xerophyllaceae Takhtajan 1996). 13/100.

Superorder LILIANAE 627

Temperate Eurasia, North America; Schoenocaulon

(24) New Mexico, Texas and Florida through

Mexico (c.20) south to Venezuela, southern Peru

(1, S. ofﬁ cinale).

xerophylleae: Xerophyllum; melanthieae:

Veratrum, Melanthium, Stenanthium, Amianthium,

Schoenocaulon, Zigadenus, Anticlea, Toxicoscordion;

heloniadeae: Helonias (including Heloniopsis –

Tanaka 1998), Ypsilandra; chionographideae:

Chamaelirium, Chionographis.

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93: 1178–1192.

Order 7. TRILLIALES

Perennial herbs with long or short (sometimes tuber-

ous), horizontal or vertical, monopodial rhizome and

mostly one erect, unbranched stem. Oxalate raphides

present in stem and leaves. Vessels only in roots, with

scalariform perforations. Sieve-element plastids of

P2c-type (Behnke 2002). Leaves in one verticil of

three (Trillium) or 4–22 (other genera) at the top of

stem, sessile or often petiolate, linear to ovate, elliptic,

rounded or rhomboid, entire, sometimes more or less

cordate at base, venation parallel to arcuate-pinnate

with more or less reticulate secondary veins. Stomata

tetracytic. Flowers always solitary and terminal,

commonly rather large, mostly on long pedicel, mostly

3-merous or 4-merous, rarely (some species of Daiswa)

5-merous, bisexual, actinomorphic, with double

perianth or sometimes apetalus. Perianth segments free.

Sepals 3–9(10), mostly green, commonly lanceolate;

petals 3–9(10), ﬁ liform to ovate, of various colours,

rarely rudimentary. Stamens 3–24 (very rarely up to

34 in some species of Daiswa), mostly 6–12 (in Trillium

usually 6), in 2–6 cycles, persistent, adnate to base of

perianth segments; ﬁ laments usually short, slender;

anthers basiﬁ xed, linear or oblong, often produced into

well-developed apical appendage (up to 12–16 mm in

Daiswa thibetica), dehiscent longitudinally, extrorse to

introrse. Tapetum secretory. Microsporogenesis suc-

cessive. Pollen grains 2-celled, 1-colpate and ellipsoi-

dal or (most species of Trillium) inaperturate and

spheroidal, tectate or intectate, reticulate, gemmate or

spinulose. Septal nectaries present. Gynoecium of 3–12

carpels (in Trillium always three); stylodia free or more

or less connate into deeply branched style that is usu-

ally separated from ovary by transverse rim; ovary

superior, roundish or angular, sometimes depressed at

the top, 3(-12) locular, 1-locular or 3(-12)-locular below

and 1-locular above, with numerous anatropous ovules

per locule or on intrusive parietal placentas. Ovules

bitegmic, crassinucellate, with parietal cell. Crystal

cells containing raphide bundles are common in the

chalaza. Female gametophyte is bisporic of Allium-type.

Endosperm helobial (Trillium) or nuclear (Paris). Fruits

more or less ﬂ eshy, berry-like or more or less capsule-

like (loculicidal in Daiswa). Seeds light to dark brown,

ellipsoidal or almost globose, smooth or reticulate-

foveolate, in Daiswa surrounded by juicy, orange or

dark-red sarcotesta and in Trillium may bear an oil-rich

elaiosome developed from the raphe and hilum; embryo

minute, globose to ovoid, undifferentiated; seed coat

has parenchymatous testa and phlobaphene containing

tegmen; endosperm copious, contains aleurone, lipids,

and starch grains. Contain steroidal saponins, ﬂ avonols

(kaempferol, quercetins), and chelidonic acid.

Alkaloids lacking, n = 5. Chromosomes usually very

large (40 μm).

Trilliales have deﬁ nite similarities with the

Melanthiales, including free stylodia, helobial

endosperm, the presence of parietal cell and crystal

raphides, as well as the combined analysis of molecu-

lar and non-molecular data (Chase, Stevenson, Wikkin

Superorder LILIANAE 629

and Rudall 1995). According to Tamura (1998),

possibly the Trilliaceae have recently been derived

from the stock of the Melanthiaceae and have greatly

changed their characteristics. However, from the

Melanthiaceae the Trilliaceae differ in basic chromo-

some number, very large chromosomes, presence of

septal nectaries, pollen morphology.

1. TRILLIACEAE

Chevallier 1827 (including Paridaceae Dumortier 1829).

5/70. Europe, Mediterranean, northeastern Anatolia,

Caucasus, Siberia, Himalayas, northeastern India, north-

ern Burma, eastern Asia, Indochina, North America.

Trillium (including Pseudotrillium), Trillidium,

Kinugasa, Daiswa, Paris.

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Paris

. Acta Phytotax. Sinica 26: 10–21.

Naumova TN. 1990. Trilliaceae. In: TB Batygina and MS Yakovlev,

eds. Comparative embryology of ﬂ owering plants:

Monocotyledons, pp. 151–159. Nauka, Leningrad (in Russian).

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Plant Spec. Biol. 4: 1–28.

Punina EO, Yu A Myakoshina, AM Eﬁ mov, and AV Rodionov.

2000. Chromosome maps of Trilliaceae plants: heterochro-

matin nucleotide composition and mapping of 18S–26S

rRNA genes in Paris quadrifolia L. Russ. J. Genet. 36:

546–549.

Smith MC and R Ingram. 1986. Heterochromatin banding in the

genus Paris. Genetica 71: 141–145.

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Takahashi M. 1982. Pollen morphology of North American spe-

cies of Trillium. Am. J. Bot. 69: 1185–1195.

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genera. Bot. Mag. (Tokyo) 97: 233–245.

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and genera of vascular plants, vol. 3, pp. 444–452. Springer,

Berlin/Heidelberg/New York.

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seed anatomy, vol. 1, pp. 130–132. Nauka, Leningrad (in

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17: 317–324.

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States. Harv. Pap. Bot. 9: 91–120.

630 Subclass II. LILIIDAE

Order 8. LILIALES

Perennial or rarely annual herbs with rhizomes (tuber-

ous in Medeola), tunicated corms or bulbs. Raphides

lacking or rarely (Campynemataceae) present. Vessels

only in roots, or both in roots and stems, with scalari-

form perforations. Leaves both cauline and basal, or

only basal, alternate or rarely verticillate, sometimes

distichous, sessile or sometimes petiolate, linear to

ovate, usually parallel-veined, often sheathing at the

base. Stomata anomocytic. Flowers small, medium-

sized, or large, in various types of inﬂ orescences or

sometimes (Tulipa spp.) solitary, bisexual or very

rarely unisexual, actinomorphic or rarely weakly zygo-

morphic (Fritillaria spp.). Perianth of six segments,

free from each other, in two cycles, generally more or

less similar or rarely (as in Nomocharis) different in

the two whorls, with nectaries on the tepal bases

(perigonal or less often androecial nectaries), or with-

out nectaries (Mediolaceae). Stamens six, in two

cycles, or rarely (Scoliopaceae) three, free from each

other or epipetalous; anthers basiﬁ xed or more often

dorsiﬁ xed, extrorse or latrorse, sometimes introrse, tet-

rasporangiate, opening longitudinally. Tapetum secre-

tory. Microsporogenesis successive. Pollen grains 2- or

3-celled, 1-colpate or rarely with 2–3 distal colpi or

inaperturate, often more or less reticulate. Gynoecium

of three united carpels; stigma on short or long style,

3-lobed or with three stigmatic crests, papillate; ovary

superior, 3-locular or sometimes 1–2-locular, with sev-

eral to numerous ovules. Ovules anatropous or rarely

campylotropous, usually bitegmic, crassinucellate or

rarely weakly crassinucellate or (in some genera of

Colchicaceae) tenuinucellate, without parietal tissue,

often with funicular obturator. The archesporial cell

functions directly as the megaspore mother cell with-

out cutting off a parietal cell. Female gametophyte tet-

rasporic, mostly of Fritillaria-type, or Polygonum-type.

Endosperm nuclear. Fruits septicidal or loculicidal

capsules, or baccate (Streptopeae and Medeola). Seeds

of various shape and size, seed coat is formed by both

integuments, but in general it lacks mechanical

strength; the testa is thin, sometimes only 2-cell-

layered; without phytomelan, but usually with phloba-

phenes; embryo mostly little differentiated; endosperm

contains aleurone and fatty oils, sometimes with traces

of starch. Flavonol glycosides based on kaempferol

and quercetin present. Steroidal saponins and steroidal

alkaloids occur in some genera.

Related to the Melanthiales.

Key to Families

1 Ovary superior.

2 Plants rhizomatous.

3 Stamens six in two cycles.

4 Fruits capsular. Herbs with usually stoloni-

form rhizome, and mostly decaying every

year, rarely thickened and surviving for a

few years. Stem erect, mostly simple.

Vessels both in roots and stem, with scalari-

form perforations which may have more

than 40 bars. Leaves alternate, sessile, all

cauline, oblong to ovate, sometimes clasp-

ing or perfoliate, reticulate-veined. Stomata

surrounded by four stomatal contact cells,

two cells lateral to the guard cells and one

at each pole. Flowers rather large, terminal

or axillary, solitary or in cymes or clusters,

bisexual, actinomorphic. Perianth segments

free to base, recurved above, caducous, the

outer ones basally provided with globose,

nectariferous pouches or less often with

short spurs, more or less purple-spotted

inside, the inner ones ﬂ at. Filaments slightly

ﬂ attened, more or less connate below,

spreading above; anthers oblong, dorsi-

ﬁ xed, extrorse. Pollen grains 1-colpate,

reticulate. Ovary superior, oblong, 3-angled,

3-locular, with numerous ovules. Stylodia

connate into a columnar style with three

spreading, bifurcate stigmas, tuberculate on

inner side. Capsules narrowly oblong to lin-

ear, many-seeded. Seeds minute, reticulate,

ovate to orbicular in outline, ﬂ at, piled upon

each other (as in several genera of Liliaceae);

testa contains lignin, tegmen totally col-

lapsed. Have large, round crystals of cal-

cium carbonate like those in Erythronium

and Lilium (Goldblatt et al. 1984). Possesses

quercetin and often kaempferol. n = 13,

rarely 12.. . . . . . . . . . . 3. tricyrtidaceae.

4 Fruits baccate. Plants with subterranean rhi-

zome-like horizontal storage stem tuber

decaying every year or short rhizome surviv-

ing for several years. Leaves cauline, in two

pseudoverticillate groups (morphologically

in dense spirals), one near the middle of

stem and consists of (5)6–9(-11) oblanceo-

late to obovate, acuminate, and basally

Superorder LILIANAE 631

attenuate leaves with three strong, conver-

gent primary veins and a large number retic-

ulate secondary veins, one at the apex

consists of 3(-5) smaller leaves of different

shape, short-petioled or sessile, mostly

ovate, acuminate, basally obtuse to more or

less truncate, with similar venation (Medeola)

or leaves crowded at the base of stem. The

basal parts of young leaves and stems are

covered with ﬂ occulent and deciduous wool

consisting of extremely long hairs that

develop from epidermal cells by mere elon-

gation. Inﬂ orescence is a terminal and ses-

sile, umbel-like cluster (synﬂ orescence) of

usually three sympodially arranged and con-

tracted monochasia (bostryxes according to

Berg 1962a) each of which is subtended by

one of the leaves of the upper whorl

(Medeola) or (Clintonia) inﬂ orescence often

a raceme, sometimes an umbel or 1-ﬂ ow-

ered. Flowers small and greenish; pedicels

deﬂ exing at anthesis and straightening after

anthesis (Medeola) or erect to horizontal

(Clintonia); perianth petaloid, with free, val-

vate segments. Nectaries lacking. Filaments

ﬁ liform; anthers small, pseudobasiﬁ xed,

extrorse. Pollen grains 1-colpate. Ovary

1- locular, with three deeply protruding pari-

etal placentas that come quite close together

(Medeola) or 2- or 3-locular (Clintonia);

style very short, deciduous, but with three

long, ﬁ liform, ﬁ nely papillate, longitudinally

grooved all along the inner side, stigmata

branches are divaricate to ﬁ nally deﬂ exed-

recurved over the anther (Medeola) or

(Clintonia) trilobate, not deciduous. Fruits

globose purplish-blue berry with 12–15

closely packed seeds. Seeds without append-

ages. Testa palisadlike. Embryo minute. n =

7 ( usually), 8.. . . . . . . . . . 7. medeolaceae.

3 Stamens three, the inner stamen cycle lacking.

Herbs with short, slender rhizome and short,

subterranean stem, which at the soil surface

bears two (rarely three or four) leaves. Leaves

alternate, distichous, elliptic to oblong, ses-

sile-clasping to subsessile, parallel veined

with numerous transverse veinless, more or

less mottled with purple spots. Inﬂ orescences

nearly sessile, 2–12-ﬂ owered, umbel-like,

consists of axillary fascicles of elongated,

3-angled, twisting or recurving pedicels; each

fascicle might be either a contracted bostryx

or a contracted cincinnus (Berg 1959) or a

contracted raceme (Utech 1992). Flowers

rather large. Perianth segments free, dissimi-

lar; outer segments petaloid, many-veined,

striped, spreading, with basal nectary glands;

inner segments linear, 3-veined, converging

over the gynoecium. Stamens opposite and

adnate basally to outer segments; ﬁ laments

free, ﬁ liform-subulate; anthers oblong,

dorsiﬁ xed, extrorse. Pollen grains 1-colpate,

spheroidal or somewhat oblong, rather large,

tectate- columellate, with rugulate arrange-

ment of ﬁ ne reticulum or foveolate. Ovary

sessile, strongly 3-angled (the dorsal regions

laterally ﬂ attened or compressed, and the ven-

tral regions occupy the corners of ovary),

1-locular, with numerous ovules on parietal

placentas; ovules with very long funicle; style

short, with three linear, horizontally spreading

to recurved stylar branches that are channeled

on the upper stigmatic surface and bear small

stigma at the apex. Fruits strongly 3-angled

and thin-walled capsule terminated by persis-

tent style with atypical irregular opening

caused by parenchymatic cell degeneration of

pericarp tissue between the six dorsal and

ventral veins in the upper part of the fruit.

Seeds oblong and slightly curved, longitudi-

nally sulcate-striate, with an appendage (elaio-

some) produced by the raphe that forms

ventral excrescence along the seed, extending

from near the micropyle to nearly across the

chalazal end; embryo minute, approximately

one tenth of the length of the endosperm,

n = 7, 8.. . . . . . . . . . . . . . . 4. scoliopaceae.

2 Plants with corm or bulb.

5 Plants with starchrich corms, in some genera

stoloniferous, somewhat (Gloriosa) rhizoma-

tous. Perennial herbs with a basal aggregation

of leaves. Vessels only in roots. Stems erect or

climbing (crimbers stem twiners and tendril

climbers). Leaves alternate, spiral (usually) or

distichous, simple, entire, linear or lanceolate,

sheathing, parallel-veined. Flowers medium-

sized to large, in bracteate raceme or spicate,

bractless inﬂ orescences, or solitary. Perianth

segments more or less similar, free from each

other or basally connate, in Colchicum and

632 Subclass II. LILIIDAE

related genera forming long tube. Nectaries

perigonal (at the bases of perianth segments)

or (Colchicoideae) androecial. Stamens

inserted at the base of segments; ﬁ laments nar-

row or basally broad; anthers mostly short,

ovate, basiﬁ xed or dorsiﬁ xed, extrorse or

latrorse, very rarely almost introrse. Pollen

grains tectate-columellate, 1-colpate or 2–4-

(-many)-porate, reticulate, foveolate-reticulate

or striate. Stylodia free or united into deeply

3-brachiate style, Ovary 3-locular, with sev-

eral to many anatropous to weakly campy-

lotropous, pseudocrassinucellate ovules per

locule. Fruits septicidal or less often loculi-

cidal capsules. Seeds globose or rarely

(Onixotis) ovoid, sometimes with more or less

well expressed raphal appendage (strophiole);

testa mostly brown or red; embryo relatively

short, elongate, or rarely subglobose; seed-

lings have conspicuously coleoptilelike coty-

ledon; endosperm oily. Lack true raphides but

have crystal sand. Most genera contain

colchicine-type (and related compounds)

alkaloids with a tropolone ring, as well as

many other alkaloids without a tropolone

ring. n = 5, 7–12, 19. . . . . 2. colchicaceae.

5 Plants with tunicate bulbs.

6 Fruits septicidal capsules. Perennial, erect,

often slender herbs with tunicated bulb hav-

ing membranous or ﬁ brous-reticulate coat.

Vessels only in roots. Leaves alternate, ﬂ at,

sessile, sheathing, simple, entire, linear to

lanceolate, few or sometimes only one basal

leaf. Flowers large, solitary or few together

in a false umbel, with long pedicels, situ-

ated in the axils of green, lanceolate, or lin-

ear bracts. Perianth differentiated into two

cycles; outer segments lanceolate to ovate,

generally glabrous, and either green and

sepaloid or similarly coloured as the inner

segments; inner segments larger and

broader, petaloid, cuneate to clawed, dis-

tinctly colored and usually marked with

large or small spots, streaks or other pat-

terns (nectar guides), more or less hairy

inside and with an often densely bearded,

depressed nectary gland or spot near the

base. Stamens six, inserted at the base of

the inner segments; ﬁ laments subulate and

basally dilated; anthers pseudobasiﬁ xed

(surrounding the tip of the ﬁ lament with a

tubular part of the connective), latrorse.

Pollen grains 1-colpate, reticulate. Style

very short or obsolete and three stigmatic

branches are situated on the apex of the

ovary; ovary 3-locular, with numerous ana-

tropous ovules per locule. Fruits 3-angled

or -winged, septicidal capsules. Seeds trig-

onous, tetragonous, or more or less irregu-

larly shaped or ﬂ attened and lenticular;

embryo slender, but well differentiated;

endosperm containing fat and aleurone.

Alkaloids sometimes present in the seeds;

n = 7–10.. . . . . . . . . .5. calochortaceae.

6 Fruits loculicidal capsules. Plants with tuni-

cated or nontunicated bulbs that have from

one (Gagea) to numerous ﬂ eshy nutrient

scales. Roots contractile. Leaves basal or

cauline, alternate or rarely verticillate, lin-

ear to ovate-lanceolate, sessile or some-

times petiolate, parallel-veined, often

sheathing at the base. Inﬂ orescence cymose

or rarely racemose and may form a raceme

or (Gagea) be umbel-like, or sometimes

reduced to single terminal ﬂ ower. Flowers

actinomorphic or rarely weakly zygomor-

phic; perianth segments similar or rarely

(Nomocharis) dissimilar. Nectaries present

at the tepal bases. Anthers epipeltate (as in

Lilium, Notholirion, and Nomocharis) when

part of the anther with its connective extends

below its point of attachment, or pseudo-

basiﬁ xed when the ﬁ lament tip is sur-

rounded by the tubular connective. Pollen

grains large, mostly monocolpate or less

often with 2–3 morphologically distal colpi,

rarely inaperturate or 1–3-porate, often

coarsely reticulate. Ovary 3- locular; style

short or long, apically 3-lobed or with three

stigmatic crests; stigmatic surface papillate;

placentas and margins of not completely

fused carpels provided with stigmatoid tis-

sue. Fruits loculicidal capsules. Seeds usu-

ally ﬂ at and disc-shaped, closely piled on

top of each other, often provided with mar-

ginal rim, sometimes with raphal elaio-

some; exotesta contain phlobaphenes,

n usually = 12. . . . . . . . . . . . 6. liliaceae.

1 Ovary inferior to semi-inferior. Slender herbs with

very short rhizome and thick adventitious roots.

Superorder LILIANAE 633

Vessels (very primitive) only in roots, but sometimes

absent. Leaves and probably also other parts contain

calcium oxalate raphide bundles in special idioblasts.

Leaves alternate, sessile, few, one basal or several

basally clustered and progressively smaller on ﬂ ow-

ering stem, linear to linear-lanceolate, sheathing at

base, in Campynemanthe basal leaf apically 3-den-

tate. Stomata anomocytic. Flowers small, in termi-

nal, bracteate few-ﬂ owered panicle, several-ﬂ owered

pseudoumbel (Campynemanthe), or solitary, bisex-

ual, actinomorphic. Perianth segments six, free,

inserted near the top of the ovary, similar, ovate or

broadly lanceolate, greenish, sometimes dotted-stri-

ated, without spots or spotted, persistent, enlarging

after fertilization, provided with perigonal nectaries

(Campynemanthe). Stamens six, inserted at base of

perianth segments; ﬁ laments slender; anthers in

Campynema sagittate and dorsiﬁ xed and extrorse,

in Campynemanthe basiﬁ xed to subbasiﬁ xed and

latrorse or weakly extrorse (Campynema). Tapetum

secretory, Microsporogenesis successive. Pollen

grains 1- colpate (Campynema) or inaperturate

(Campynemanthe), reticulate. Gynoecium of three

united carpels; stylodia free (Campynema) or con-

nate into apically 3-lobed style (Campynemanthe);

ovary inferior or semi- inferior (Campynemanthe),

3- locular, or 1-locular (Campynema), with few to

many ovules; placentation when unilocular parietal,

when trilocular –axile. Ovules anatropous, bitegmic,

weakly crassinucellate. The parietal cell is formed

from the archesporial cell and gives rise to parietal

tissue. Fruits capsules dehiscing by decay of the

lateral or dorsal walls or indehiscent. Seeds pale red-

brown, in Campynemanthe nearly globose to angular

and with long persistent funicles, in Campynema

ﬂ attened and with spongy outer coat; seed coat

formed (at least in Campynemanthe) by both integu-

ments and both the testal and tegmic parts are 2-celled

layer thick; the endotegmic layer contains phloba-

phene; embryo minute; endosperm copious. Contain

saponins; n = 11.. . . . . . . . . .1. campynemataceae.

1. CAMPYNEMATACEAE

Dumortier 1829. 2/4. New Caledonia (Campynemanthe)

and Tasmania (Campynema).

Campynemanthe, Campynema.

The afﬁ nities of this rather isolated family is uncer-

tain. Although it is often connected with or even

included in the Melanthiaceae s. l., the nuclear devel-

opment of endosperm (Lowry et al. 1987) makes it

closer to the colchicalean families. However,

Campynemataceae markedly differ from all other fam-

ilies of the Liliales by the presence of crystal raphides

and almost inferior ovary.

2. COLCHICACEAE

A.P. de Candolle 1804 (including Burchardiaceae

Takhtajan 1996; Merenderaceae Mirbel 1804;

Uvulariaceae A. Gray ex Kunth 1843). 23/95. Africa

(especially South Africa), Eurasia, Australia, New

Zealand.

2.1 UVULARIOIDEAE

Plants with creeping or short rhizome. Stamens six, free;

ﬁ laments expanded at base; anthers, extrorse or

(Kuntheria) introrse. Pollen grains 1-colpate or

(Uvularia) usually 2-colpate, tectate-columellate,

foveolate-verrucate. Fruits somewhat ﬂ eshy loculi-

cidal capsules or (Uvularia) septicidal capsules or

(Streptopeae) baccate. Seeds without elaiosome. Contain

calcium oxalate in the form of raphides, sometimes also

crystal sand. Colchicin absent. n = 6, 7, 8, 9. –

uvulareae: Uvularia, Schelhammera, Tripladenia,

Kuntheria; streptopeae: Prosartes, Disporum,

Streptopus

; burchardieae: Burchardia.

2.2 COLCHICOIDEAE

Plants with starch corms, in some genera stolonifer-

ous. Stamens inserted at the base of perianth seg-

ments. Nectaries perigonal (at the bases of perianth

segments) or androecial (Colchiceae) Pollen grains

1-colpate 2- or 4- to many-porate, reticulate, or stri-

ate. Most genera contain colchicines-type alkaloids.

– wurmbeeae (anguillarieae): Neodregea,

Onixotis, Wurmbea, Baeometra; iphigenieae:

Iphigenia, Camptorrhiza, Ornithoglossum,

Hexacyrtis; glorioseae: Gloriosa, Littonia,

Sandersonia; colchiceae: Androcymbium,

Merendera, Bulbocodium, Colchicum.

3. TRICYRTIDACEAE

Takhtajan 1997 (including Compsoaceae Horaninow

1834). 1/18. Eastern Himalayas, northeastern India

(Khasia Hills), China, Taiwan, Japan.

634 Subclass II. LILIIDAE

Tricyrtis.

Both chemical evidence (Williams et al. 1988) and

morphological data support the separation of Tricyrtis

from the Colchicaceae where it is usually placed.

4. SCOLIOPACEAE

Takhtajan 1996. 1/2. Western North America along the

Paciﬁ c coast from Santa Cruz Mountains in California

to the Cascade Mountains in Oregon.

Scoliopus.

The taxonomic position of the genus Scoliopus has

been debated for a long time. At different times it has

been included in Melanthiaceae s. l., Colchicaceae,

Medeoleae of the Liliaceae s. 1., and Trilliaceae.

Dahlgren et al. (1985: 231) concludes that Scoliopus

“is perhaps best placed in Uvulariaceae.” Takhtajan

(1987) placed the genus in a separate tribe Scoliopeae

within Melanthiaceae-Melanthioideae, Tamura (1998)

in Calochortaceae-Tricyrtidoideae, and Thorne (2006)

includes it in Liliaceae. However, Scoliopus differs

markedly from both the Melanthiaceae s. str.,

Uvulariaceae and Calochortaceae in many important

features, including peculiar vegetative morphology,

inﬂ orescence, reduced androecium, 1-locular ovary

with nonintruding parietal placenta, unique type of the

fruit dehiscence, and seeds provided with strophiole.

5. CALOCHORTACEAE

Dumortier 1829. 1/65. Temperate western North

America from British Columbia and Dakota south to

Guatemala with the greatest concentration of species

in California.

Calochortus.

Calochortaceae are rather isolated within the order,

but embryologically they are nearest to the

Scoliopaceae (“The greatest embryological similarity

is found between Scoliopus and Calochortus,” Berg

[1962: 51]).

6. LILIACEAE

A.L. de Jussieu 1789. (Erythroniaceae Martynov,

1820; Fritillariaceae Salisbury, 1866; Liriaceae Batsch

ex Borkhausen 1797; Tulipaceae Batsch ex Borkhausen

1797). 13/550. Temperate and subtropical regions of

the Northern Hemisphere, especially western, Central

Asia, Himalayas, and eastern Asia.

lloydieae: Gagea, Lloydia; lilieae: Cardiocrinum,

Lilium, Notholirion, Nomocharis, Fritillaria Korol-

kowia, Rhinopetalum; tulipeae: Tulipa, Amana,

Holongia, Erythronium.

7. MEDEOLACEAE

Takhtajan 1987. 2/6. Eastern Asia and North

America.

clintonieae: Clintonia; medeoleae: Medeola.

Related to the Liliaceae but differ from them

markedly in basic chromosome number, type of the

subterranean storage organ, the absence of nectaries,

style morphology, crassinucellate ovules, and baccate

fruits.

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