Takhtajan Armen. Flowering Plants

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Superorder LILIANAE 665

quantities of starch (more abundant at

the early stages of development); embryo

straight or slightly curved, generally

more than half as long as the endosperm.

Steroidal saponins lacking, but various

particular alkaloids as well as chelidonic

acid and norbelladine alkaloids are

widely distributed. n = 5–15, 23, 27, 29,

mostly 11.. . . . . .13. amaryllidaceae.

10 Ovary superior. Plants rhizomatous.

Roots ﬂ eshy, provided with a multiple

velamen. Laticifers present. Vessels only

in roots, mostly with scalariform perfora-

tions. Calcium oxalate raphides present.

Leaves rosulate, distichous, linear, sheat-

ing at base. Flowers in a pseudo-umbel

subtended by two involucral spathial

bract, bisexual, zygomorphic, pedicels

subtended by threadlike bracts. Perianth

segments subequal, basally more or less

connate; corona lacking. Stamens

inserted on the perianth tube; ﬁ laments of

unequal length, declinate; anthers dorsi-

ﬁ xed. Pollen grains 1-colpate, reticulate.

Style slender, hollow, declinate, with

punctiform stigma; ovary 3-locular, with

inner septal nectaries. Ovules numerous,

campylotropous, crassinucellate. Parietal

cell present. Fruits loculicidal capsules.

Seeds black, angulate, elongated, winged;

endosperm stores aleurone, lipids and

reserve cellulose; embryo well devel-

oped, 4/5 of the length of endosperm

Various steroidal saponins present. n =

15, rarely 16. . . . 11. agapanthaceae.

11 Inﬂ orescences monopodial racemes

or spike. Rootstock mostly a bulb.

Perennial herbs with bulbs or rarely

(Schoenolirion and Chlorogalum)

rhizomes; bulbs generally with mem-

branous tunic and free or coalescent

scales. Roots sometimes thick and

commonly contractile. Vessels only

in roots, with scalariform and/or sim-

ple perforations. Leaves concentrated

at the base, linear to lanceolate, rarely

elliptic to orbicular, sessile, sheathing

at base. Inﬂ orescences monopodial

raceme or spike. Flowers not articu-

lated, generally bisexual, actinomor-

phic or rarely zygomorphic. Perianth

segments free or more often connate.

Stamens six in two cycles, with some-

times one sterile cycle; ﬁ laments

inserted at base of perianth segments

or on the perianth tube; often broad

and ﬂ at; sometimes connate at the

base, rarely forming a paracorolla;

anthers epipeltate. Pollen grains,

1-colpate. Style simple, with puncti-

form or sometimes distinctly 3-lobed

stigma; ovary superior or rarely

(Bowiea) semi- inferior, 3-locular,

with septal nectaries (except for

Autonoe), with two to numerous ana-

tropous or campylotropous ovules

per locule. Female gametophyte of

various types, but mostly of

Polygonum-type. Endosperm helo-

bial or nuclear. Fruits loculicidal cap-

sules. Seeds small to large, ovoid to

obovate, with elaiosomes; outer epi-

dermis of the testa consists of several

layers and usually has thick to rather

thin phytomelan crust (absent in

Chionodoxa, Pushkinia, and a few

species of Scilla, where outer epider-

mis collapses); inner integument

forms thin membrane; endosperm

usually copious, consists of pitted

cells and contains aleurone and fatty

oils and very rarely starch grains;

embryo more or less cylindrical, usu-

ally straight. Crystal raphides con-

tained in mucilage cells or canals are

widely distributed. Producing steroi-

dal saponins, chelidonic acid, poly-

hydroxyalkaloids, ﬂ avones, ﬂ avone

C-glycosides. n = 2–15, 17, 19, 21,

27, 29. . . . . . . . . 8. hyacinthaceae.

11 Inﬂ orescences contracted cymes,

rarely spikelike or umbellate.

12 Rootstock a bulb. Leaves spiral.

Corona present. Alliaceous odour

usually present. Rootstock bulbs

or bulb-like corms with membra-

nous or ﬁ brous scales, more rarely

rhizome. Articulated laticifers

666 Subclass II. LILIIDAE

present in leaves and bulb scales.

Vessels only in roots, with scalari-

form or/and simple perforations.

Leaves basally concentrated, ﬁ li-

form-linear, lanceolate or rarely

ovate, sheathing at base, rarely

constricted into a pseudo petiole.

Flowers usually in one or more

contracted cymes, rarely (as in

Milula) spikelike, subtended by

involucre of (1) 2 or more mem-

branous spathial bracts, free or

basally united, generally bisex-

ual, actinomorphic or zygomor-

phic, not articulated. Perianth

segments free or often connate.

Functional stamens six or some-

times three or two with several

staminodia; ﬁ laments inserted at

the base of perianth segments or

in perianth tube, more or less

ﬂ at; anthers mostly elongate,

epipeltate. Pollen grains, 1-col-

pate. Style simple, with 3-lobed

or capitate stigma; ovary superior,

3-locular, with two to several ana-

tropous, hemitropous, or campy-

lotropous, crassinucellate (as in

Milula) or tenuinucellate (as in

Allium, Nothoscordum,

Tulbaghia) ovules. Septal necta-

ries present. Female gametophyte

of Polygonum- or Allium-type.

Endosperm helobial. Fruits locu-

licidal capsules. Seeds of various

sizes and shapes, with testal epi-

dermis usually covered by rather

thick crust of phytomelan and

inner layers of testa compressed

or collapsed, as is the tegmen;

endosperm cells with rather thick

and pitted walls and contain aleu-

rone and fatty oils; embryo

straight or more or less curved

and usually more than half the

length of endosperm. Contain ﬂ a-

vonoids, steroidal saponins and

strong-smelling allyl disulﬁ des,

propyl sulﬁ des and vinyl sulﬁ des

that participate in essential oils

causing the onion odour. n = 5–16.

. . . . . . . . . . . . . 12. alliaceae.

12 Rootstock a starch-storing corm;

a new corm is forming every year

on top of the corm of the previous

year. Lacticifers present. Calcium

oxalate raphides present or absent.

Vessels with scalariform and sim-

ple perforations. Leaves spirally

arranged, linear, ﬂ at, forming

closed sheath. Flowers in umbel-

late inﬂ orescence, rarely reduced

to single ﬂ ower; pedicels often

articulated, subtended by a small

bract (two bracts when ﬂ ower is

solitary). Perianth segments usu-

ally more or less basally connate.

A corona, scales or appendices

often present between perianth

segments and stamens. Fertile

stamens three, the missing ones

often transformed into stamino-

dia; anthers more or less bas-

iﬁ xed, introrse. Pollen grains

1-colpate, reticulate. Style with

3-lobed or capitate stigma; ovary

sometimes on a gynophore, 3-loc-

ular, with two or several anatro-

pous ovules per locule. Endosperm

helobial (Muilla and Triteleia) or

nuclear (Brodiaea). Fruits loculi-

cidal capsules. Seeds angular;

epidermis of testa with a crust of

phytomelan; embryo short and

thick; endosperm copious, con-

taining aleurone and lipids.

Alliaceous odour lacking.

Steroidal saponins present or

absent. n = 5–12, 14, 15, 21, 25,

27.. . . . . . . . . 10. themidaceae.

1. HEMEROCALLIDACEAE

R. Brown 1810. 1/ about 15. Central Europe (1) to

Japan and China, mainly in China.

Hemerocallis.

The least advanced member of the order.

Superorder LILIANAE 667

2. PHORMIACEAE

J. Agardh 1858. (including Dianellaceae R.A. Salisbury

1866). 10/30–35. Mediterranean (Simethis) Mada gascar,

Mascarenes, Seychelles, tropical East Africa, southern,

eastern and southeastern Asia, Malesia, Hawaii, South

Paciﬁ c Islands, Fiji, New Caledonia, New Zealand,

Australia, Chile (Pasithea), and one monotypic genus

Excremis in Andes of Ecuador, Colombia and Peru;

Phormium – New Zealand (N and S Isl., Stewart Isl.,

Chatham Isl., Auckland Isl.) and Norfolk Island.

2.1 PHORMIOIDEAE

Rhizome contains wax and leaves contain

hemicellulose-like polysaccharide. Raphides absent.

Seeds winged. Proanthocyanidins absent. – Phormium.

2.2 DIANELLOIDEAE

Wax in rhizome and hemicellulose-like poly-saccha-

rides in leaves absent. Raphides present Seeds wingless.

Proanthocyanidins present (Dianella). – Agrostocrinum,

Pasithea, Herpolirion, Thelionema, Simethis, Stypandra,

Rhuacophila, Dianella, Excremis.

Related to the Hemerocallidaceae.

3. XERONEMATACEAE

M.W. Chase, Rudall and Fay, 2000. 1/2. New Caledonia,

New Zealand.

Xeronema.

Possibly related to the Phormiaceae.

4. ASPHODELACEAE

A.L. de Jussieu 1789 (including Aloaceae Batsch

1802). 15/1000. Temperate, subtropical regions of the

Old World, Alooideae – Africa (especially South

Africa), Madagascar, Mascarenes, Arabia.

4.1 ASPHODELOIDEAE

Leaves generally not conspicuously succulent. Stomata

generally anomocytic. Vessels with scalariform perfora-

tions, only occasionally present in the stem (species of

Asphodelus). Typical caps of aloine cells are not formed

at the phloem poles of the vascular bundles of leaves.

Herbaceous plants. n = 6–11 – asphodeleae: Eremurus,

Asphodelus, Asphodeline, Bulbine, Bulbinella, Jodrellia,

Trachyandra, Paradisea; kniphofieae: Kniphoﬁ a.

4.2 ALOOIDEAE

Leaves more or less strongly succulent, often prickly

along the margins. Stomata mostly sunken, more or

less distinctly tetracytic. Large cap of aloine cells

present at the phloem pole of most vascular bundles

of leaves. Sparsely branched trees up to several meters

tall to small rosette herbs, n mostly = 7, with four

long and three short chromosomes. – Aloe, Poellnitzia,

Lomatophyllum, Gasteria, Haworthia, Chortolirion.

5. XANTHORRHOEACEAE

Dumortier 1829. 1/30. Australia and Tasmania.

Xanthorroea.

6. ANTHERICACEAE

J. Agardh 1858 (including Boryaceae Rudall, Chase et

Conran 1997; Johnsoniaceae Lotsy 1911;

Laxmanniaceae Bubani 1902). 27/c.350. Africa,

Madagascar, Mascarenes, Eurasia, Malesia, Australia,

Paciﬁ c Islands, North, Central and South America.

The largest genera is Chlorophytum (about 300).

anthericieae: Anthericum, Comospermum,

Chlorophytum, Dichopogon, Hagenbachia, Diamena,

Diora, Eremocrinum, Leucocrinum, Echeandia; john-

sonieae: Tricoryne, Hodgsoniola, Caesia, Corynotheca,

Arnocrinum, Hensmania, Johnsonia, Stawellia; lax-

mannieae: Arthropodium, Murchisonia, Thysanotus,

Trichopetalum, Laxmannia, Sowerbaea, Chamaescilla;

boryeae: Alania, Borya.

7. ANEMARRHENACEAE

Conran, MW Chase and Rudall 1997. 1/65. Northern

China and Korea.

Anemarrhena

8. HYACINTHACEAE

Batsch ex Borkhausen 1797 (Eucomidaceae Salisbury

1866; Lachenaliaceae Salisbury 1866; Ornithogalaceae

Salisbury 1866; Scillaceae Vest 1818). 67/800–1000.

Very widely distributed in both hemispheres, but espe-

cially diversiﬁ ed in South Africa, Mediterranean, and

Irano-Turanian region.

668 Subclass II. LILIIDAE

8.1 OZIROËOIDEAE

Bulb leaves imbricate, mucilaginous. Foliage leaves

few, thick and grooved. Scape with prominent bracts,

1–2(-3) ﬂ owered. Pedicels straight, pointing obliquely

upwards. Stamens basally connate and adnate to the

corolla. Seeds rounded; embryo as long as seed;

n = 15, 17. – Oziroë.

8.2 ORNITHOGALOIDEAE

Bulb leaves usually lasting only 1–2 vegetation peri-

ods. Bracts large, prophylls lacking. Stamens three;

ﬁ laments ﬂ at, often appendaged. Seeds often com-

pressed and edged, ridged, or carinate. Cardenolides

partly present; n = 2–10. – dipcadieae: Stellarioides,

Coilonox, Pseudogaltonia, Dipcadi, Galtonia,

Zahariadia, Melomphis, Cathissa, Eliokarmos,

Loncomelos, Honorius; ornithogaleae: Ornitho-

galum, Albuca.

8.3 URGINEOIDEAE

Bulb leaves usually imbricate. Scapes usually solitary,

bracts spurred. Filaments not appendaged. Seeds

ﬂ attened, sometimes winged; testa brittle, not tightly

adherent to endosperm. Bufodienolids present; n = 6,

7, 10. – Bowiea, Schizobasis, Igidia, Urgineopsis,

Rhadamanthus, Litanthus, Rhadamanthopsis,

Thuranthos, Tenicroa (including Sypharissa), Drimia

Urginea.

8.4 HYACINTHOIDEAE

Bulb leaves lasting 2–3 vegetation periods. Bracts usually

small, rarely large or lacking. Prophylls present or

absent. Seeds globose, drop-shaped or ellipsoid;

elaiosomes formed from various parts of the testa;

nucleus lacking protein crystals. Homoisoﬂ avones

present; n = 4–10. – pseudoprospereae:

Pseudoprospero; massonieae: Ledebouria, Resnova,

Drimiopsis, Lachenalia, Veltheimia, Eucomis,

Massonia, Namophila, Whiteheadia, Periboea, Polyxena,

Androsiphon, Amphisiphon, Neobakeria, Daubenya;

hyacintheae: Hyacinthus, Merwilla, Schizocarpus,

Fortunatia, Barnardia, Oncostema, Hyacynthoides,

Chionodoxa, Puschkinia, Hyacinthella, Brimeura,

Tractema, Alrawia, Prospero, Othocallis, Pfosseria,

Nectaroscilla, Bellevalia, Muscari, Muscarimia,

Leopoldia, Pseudomuscari, Scilla, Rhodocodon,

Daubenya, Neopatersonia.

Close related to the Anthericaceae and Agavaceae.

9. AGAVACEAE

Dumortier 1829 (including Chlorogalaceae Doweld et

Reveal 2005; Funkiaceae Horaninow 1834;

Hesperocallidaceae Traub 1972; Hostaceae Mathew

1988; Yuccaceae J. Agardh 1858). 15/300. Tropical

and subtropical regions of America and West Indies,

concentrated mostly in Mexico and southwestern and

southern United States; primarily in dry areas.

9.1 CHLOROGALOIDEAE

Bulb leaves usually present, short-lived; scale and foli-

age leaves with little mucilage. Ovary superior, with

septal nectaries; nectar ducts tubular or covered by

hairs. – Schoenolirion, Hastingsia, Camassia,

Chlorogalum.

9.2 YUCCOIDEAE

Caespitose or abrorescent plants. Ovary superior. –

Yucca, Hesperaloe.

9.3 AGAVOIDEAE

Ovary inferior. – Agave, Beschorneria, Furcraea,

Littaea, Manfreda, Polianthes, Prochnyanthes.

9.4 HESPEROCALLIDOIDEAE

Rootstock bulbous (Hesperocallis) or not bulbous

(Hosta). Ovary superior. – Hosta. Hesperocallis.

10. THEMIDACEAE

R.A. Salisbury 1866. 12/60. Southwestern Canada,

western United States and Mexico, with one species of

Milla reaching Guatemala.

Muilla, Bloomeria, Dandya, Milla, Bessera,

Petronymphe, Triteleia, Jaimehintona, Triteleiopsis,

Brodiaea (Themis), Dichelostemma, Androstephium.

Related to the Hyacinthaceae and Alliaceae.

11. AGAPANTHACEAE

Voigt 1850. 1/9. South Africa from the Cape Peninsula

to the south of Limpopo River.

Agapanthus.

Superorder LILIANAE 669

12. ALLIACEAE

Borkhausen 1797 (including Cepaceae Salisbury 1866;

Gilliesiaceae Lindley 1826; Milulaceae Traub 1972;

Tulbaghiaceae Salisbury 1866). 18/650. Widely dis-

tributed in both hemispheres except for tropical

regions, Australia, and New Zealand.

12.1 TULBAGHIOIDEAE

Plant often rhizomatous or bulb with thick roots and

alliaceous smell; rhizomes or bulb containing starch.

The closed sheaths at base of leaves very short. Flowers

bracteate, in umbellate inﬂ orescences. Perianth seg-

ments strongly connate, corona massive, lobes connate

or not. Embryo straight or slightly curved, phytomelan

crust rather thin; n = 6. – Tulbaghia.

12.2 ALLIOIDEAE

Mostly with tunicated bulbs and with alliaceous odour;

bulbs lacking starch. Leaves more or less unifacial.

Corolla basally connate, without corona. Stamens both

basally connate and adnate to the corolla, often with

winged ﬁ laments. Embryo curved; n = (7) 8 (9).

Allium, Caloscordum, Nectaroscordum, Milula,

Nothoscordum, Ipheion, Tristagma, Garaventia.

12.3 GILLIESIOIDEAE

Corona absent or of small scales. Stamens 2–3, vari-

ously connate and adnate; staminodes present. Embryo

short; n = 6, 7, 9–10, 12. Schickendantziella, Speea,

Leucocoryne, Latace, Miersia, Gilliesia, Solaria

(including Ancrumia, Gethyum), Erinna, Trichlora.

Close to the Hyacinthaceae. According to Dahlgren

et al. (1985), most of characters found in Alliaceae are

found in the Hyacinthaceae although its inﬂ orescence

is racemose and a parietal cell is cut off in the nucellus,

a condition rare in Alliaceae.

13. AMARYLLIDACEAE

J. Saint-Hilaire 1805 (incluing Brunsvigiaceae

Horaninow 1834; Crinaceae Vest 1818; Cyrtanthaceae

Salisbury 1866; Galanthaceae G. Meyer 1836;

Gethyllidaceae Raﬁ nesque 1838; Haemanthaceae

Salisbury 1866; Leucojaceae Batsch ex Borkhausen

1797; Narcissaceae A.L. de Jussieu 1789; Oporantha-

ceae Salisbury 1866; Pancratiaceae Horaninow 1847;

Strumariaceae Salisbury 1866; Zephyranthaceae

Salisbury 1866). 61/850. Tropics and subtropics,

especially America, Mediterranean, and Africa.

Classiﬁ cation after A.W. Meerow and D.A. Snijman

(1998) and A.W. Meerow et al. (2000, 2002).

amaryllideae: Amaryllis, Boophone, Crinum,

Ammocharis, Cybistetes, Crossyne, Strumaria

(including Bokkeveldia, Gemmaria and Tedingea),

Carpolyza, Nerine, Hessea, Namaquanula,

Brunsvigia; cyrtantheae: Cyrtanthus; hae-

mantheae: Clivia, Cryptostephanus, Scadoxus,

Haemanthus; calostemmateae: Calostemma,

Proiphys; gethyllideae: Apodolirion, Gethyllis;

lycori deae: Lycoris, Ungernia; pancratieae:

Pancratium (including Chapmanolirion); narcis-

seae: Narcissus, Braxireon, Sternbergia; gal-

antheae: Galanthus, Leucojum, Vagaria, Lapiedra,

Hannonia; hippeastreae: Hippeastrum, Worsleya,

Grifﬁ nia, Rhodophiala, Zephyranthes (including

Cooperia), Habranthus, Sprekelia, Pyrolirion,

Placea, Traubia, Phycella; eucharideae

: Eucharis,

Caliphruria, Plagiolirion, Urceolina; hymenocal-

lideae: Hymenocallis, Leptochiton, Pamianthe,

Ismene; stenomesseae: Stenomesson, Phaedranassa,

Rauhia, Eucrosia, Mathieua; clinantheae:

Pseudostenomesson, Paramongaia, Pucara; euste-

phieae: Chlidanthus, Castellanoa, Eustephia,

Hieronymiella.

Related to the Alliaceae and differing from them

mainly in their inferior ovary. Although they differ

from the Alliaceae also in their unique alkaloids and

in not containing steroidal saponins, Amaryllidaceae

contain the same kind of mannose-speciﬁ c lectins,

which have similar molecular structures, agglutina-

tion properties, and amino acid composition and are

serologically identical (Van Damme et al. 1991). The

monophyly of Alliaceae and Amaryllidaceae is sup-

ported by rbcL sequences (Fay et al. 1994). It is inter-

esting to note that in Amaryllidaceae plastids form are

P2c, except Rauhia and Worsleya, which contain

form-P2cf.

14. HERRERIACEAE

Endlicher 1841. 3/12. Temperate and subtropical

South America (Herreria, Clara) and Madagascar

(Herreriopsis).

Herreria, Clara, Herreriopsis.

Probably related to the Agavaceae.

670 Subclass II. LILIIDAE

15. APHYLLANTHACEAE

Burnett 1835. 1/1. Western Mediterranean.

Aphyllanthes

A wide variety of families have been proposed

for this taxonomically rather isolated taxon, inclu-

ding Eriocaulaceae, Xanthorrhoeaceae, Liliaceae-

Asphodeloideae, and Anthericaceae. Aphyllanthes

shares a number of features with the Anthericaceae

(including septal nectaries, helobial endosperm, seeds

with phytomelan, and steroidal saponins).

Afﬁ nity is not clear, but probably belong to the

Amaryllidales.

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Order 15. ASPARAGALES

Perennial rhizomatous or tuberous (Eriospermaceae

and some Asparagaceae) herbs or less often secondary

arborescent plants often with extensive secondary

thickening growth. Oxalate raphides generally present.

Vessels only in roots or also in leaves and/or stem with

scalariform and/or simple perforations. Leaves both

basal and cauline or only basal, sessile or sometimes

petiolate, alternate, occasionally distichous or tristic-

hous, linear to ovate or cordate or reduced and scale-

like, usually parallel-veined, often scleromorphic or

succulent. Stomata anomocytic or less often paracytic

or tetracytic. Flowers in various kinds of inﬂ orescences,

sometimes solitary, bisexual or less often unisexual,

generally actinomorphic, usually 3-merous. Perianth

usually of six segments in two cycles, similar or less

often dissimilar, free or more or less connate. Stamens

usually six, rarely four or eight, in two cycles; ﬁ laments

linear, inserted at the base of perianth segments or the

perianth tube, free from each other or connate into a

column (Ruscaceae) or only basally (Peliosanthes);

anthers tetrasporangiate, ovate to elongate, basiﬁ xed or

dorsiﬁ xed, introrse or extrorse, opening longitudinally.

Tapetum secretory. Microsporogenesis successive.

Pollen grains 2-celled or rarely 3-celled, 1-colpate,

rarely inaperturate. Gynoecium of three united carpels;

style short or long, with 3-lobed, capitate or puncti-

form stigma; ovary superior or sometimes semi-inferior

or inferior, 3-locular to less often 1-locular, rarely

(some Convallariaceae) 4-or 2-locular, with 1–2 to

many ovules per carpel. Septal nectaries mostly pres-

ent. Ovules anatropous, hemitropous, campylotropous,

or sometimes orthotropous, bitegmic, crassinucellate

or tenuinucellate, mostly with a parietal cell. Female

gametophyte of Polygonum-type or sometimes of

Allium-type. Endosperm helobial or nuclear. Fruits

capsular, baccate, or dry and indehiscent. Seeds of

various sizes, shapes, and colours, with or without

phytomelan; seed coat formed mainly by the outer