Takhtajan Armen. Flowering Plants

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Superorder LILIANAE 645

Key to families

1 Ovary superior.

2 Seeds brown, clothed with colourless hairs often

appearing white. Perennial herbs with more or

less tuberous, short rhizome and ﬁ brous roots.

Leaves alternate, distichous, usually concen-

trated at the base of stem, linear, with prominent

midrib and rough edges, sheathing slightly at

base; cauline leaves small, bracteal. Stomata

anomocytic. Flowers large, pendulous, in termi-

nal racemes, bisexual, actinomorphic, campanu-

late, each subtended by a bract and two smaller

bracteoles. Perianth segments six, petaloid, con-

nate into a tube for about four-ﬁ fths of their

length. Stamens six in two cycles, inserted in

perianth tube; anthers dorsiﬁ xed, tetrasporan-

giate, dehiscing latrorsely by slits. Tapetum

secretory. Microsporogenesis simultaneous.

Pollen grains 1-colpate, with granular exine.

Ovary superior, 3-locular, tapering below into a

gynophore, stipitate, with 40–50 ovules per loc-

ule, inserted in two rows; style short, not distinct

from ovary, with small 3-grooved stigma. Ovules

anatropous, bitegmic, crassinucellate. Septal

nectaries present as deep exterior grooves.

Female gametophyte of Polygonum-type.

Endosperm helobial. Fruits septicidal capsules.

Seeds 5–6 mm long, oblong-falcate, angulate,

with phytomelan in testa and phlobaphene in

tegmen, with remnants of nucellar tissue;

endosperm copious; embryo linear, slightly

curved. n = 17, 27. . . . . . 1. blandfordiaceae.

2 Seeds black, glabrous. Perennial herbs (dwarf to

large), shrubs or trees (Cordyline) to 10 m tall,

often rhizomatous or tuberous, occasionally epi-

phytic (on trees). Raphides present. Secondary

thickening anomalous (Cordyline) or absent.

Vessels only in roots, with scalariform perfora-

tions. Leaves alternate, spiral or distichous,

mesomorphic to leathery and stiff, linear to

broadly lanceolate, mostly keeled, sheathing at

base, usually densely pubescent, at least when

young; the trichomes scalelike or stellate, multi-

cellular and arising from multicellular base;

mucilage canals present in leaves of some spe-

cies of Astelia and Collospermum. Stomata para-

cytic. Flowers in one terminal and several lateral

bracteate racemes or spikes, trimerous or

(Neoastelia) penta- to heptamerous, bisexual,

polygamo-dioecious or dioecious, actinomor-

phic. Perianth segments six in two cycles, free or

basally connate, similar. Stamens six in two

cycles, free or inserted on the perianth in

Milligania; anthers basiﬁ xed to dorsiﬁ xed, open-

ing longitudinally, introrse, unappendaged.

Female ﬂ owers with staminodia. Floral nectaries

present Microsporogenesis successive, or

(Milligania) simultaneous. Pollen grains 1-col-

pate or (Collospermum) both 1-colpate and tri-

chotomocolpate, more or less echinate.

Gynoecium of 3(-7) united carpels; stylodia con-

nate into short style (in Neolitsea stylar branches

elongated with decurrent stigmas) or stigmas

more or less sessile; in some species of Astelia

there is a gradual transition between ovary and

style. Ovary superior, 3-locular (placentation

axile) or 1- locular (placentation parietal), with

four to many anatropous ovules per locule; muci-

lage secreting intraovarian trichomes often pres-

ent; septal nectaries present. Ovules, anatropous

or campylotropous, bitegmic, crassinucellate.

Fruits berries or (Milligania) loculicidal capsules

splitting from the apex. Seeds ovate and often

angular, black, more or less shiny, with mucilagi-

nous hairs; exotesta encrusted with phytomelan;

endosperm copious, containing aleurone and lip-

ids; embryo straight or slightly bent, one third to

half the length of endosperm. Saponins/sapogen-

ins present; n = 8, 30, 35? . . . . .2. asteliaceae.

1 Ovary inferior

3 Stamens and style fused into gynostemium.

Strongly mycotrophic or occasionally sapro-

phytic perennial herbs, terrestrial or more often

epiphytic, sometimes climbing or scandent, very

rarely completely subterranean or rheophytic.

Terrestrial species with ﬁ brous or thickened

tuberous or cordlike roots and usually rhizoma-

tous, the epiphytic species often with one or sev-

eral internodes modiﬁ ed into water storage

organs (pseudobulbs) and often with aerial hang-

ing roots covered by a thick layer of dead and

usually spongy water-absorbing tissue derived

from epidermis (velamen) that are also present in

most terrestrials. Stems sympodial or (in certain

Orchidoideae) monopodial. Silica bodies

enclosed in special cells, calcium oxalate raphi-

des, and mucilage cells generally present. Vessels

mostly in roots, less often also in stem, very

646 Subclass II. LILIIDAE

rarely also in leaves, mostly with scalariform per-

forations. Leaves alternate, mostly distichous,

rarely opposite or verticillate, simple and usually

entire, sometimes plicate, linear to orbicular,

sometimes pseudopetiolate, usually parallel-

veined, often ﬂ eshy, sheathing basally with the

sheaths generally closed and enveloping the stem,

in saprophytic species reduced into small scales

or rarely entirely absent. Stomata paracytic, less

often anomocytic, sometimes tetracytic. Flowers

in racemes, spikes, or racemose panicles, some-

times solitary, bracteate, bisexual or rarely uni-

sexual (monoecious or dioecious), usually strongly

zygomorphic, 3-merous, mostly resupinate at

anthesis. Perianth segments six in two cycles;

sepals green or more often coloured and petaloid,

all similar or the median one (morphologically

abaxial) differing in size and colour, mostly imbri-

cate; the median petal (morphologically adaxial)

commonly much larger, often highly modiﬁ ed

and differing in colour from laterals forming the

labellum, which often projects basally into a spur

or sac with or without nectar, the laterals usually

more or less like sepals. Stamens three

(Apostasioideae), two (Cypripedioideae), or more

often only one (Orchidoideae), all morphologi-

cally abaxial (but apparently adaxial), opposite

labellum, when one united with the style forming

gynostemium; anthers tetrasporangiate, opening

longitudinally, introrse. Tapetum secretory.

Microsporogenesis simultaneous. Pollen grains

2-celled, solitary (Apostasioideae and part of

Cypripedioideae), in tetrads, or agglutinated into

waxy or mealy masses (pollinia), tectate-columel-

late, or sometimes tectate-granular or even exine-

less, 1-colpate, 2-colpate, 1-porate or inaperturate,

rarely (Vanilla) 3-porate, with different types of

ornamentation. Gynoecium of three united car-

pels; style more or less apically inﬂ exed and ter-

minated by a 3-lobed stigma; all three stigma

lobes may be fertile, but most commonly the two

lateral ones are fertile and the median lobe is

modiﬁ ed into a small sterile outgrowth, the rostel-

lum; ovary inferior, 3-locular or generally 1-locu-

lar, with very numerous and minute ovules per

locule or per parietal placenta. Ovules anatropous,

usually bitegmic, tenuinucellate, without parietal

cell. Female gametophyte of Polygonum-type or

(mainly Cypripedioideae) of Allium-type, very

rarely tetrasporic. Endosperm, when it develops,

is nuclear, but usually endosperm formation is

arrested very early or wholly omitted. Fruits usu-

ally capsules dehiscing by 3(6) hygroscopically

sensitive valves, rarely ﬂ eshy and indehiscent.

Seeds extremely numerous, commonly minute,

mostly dustlike, often fusiform, usually germinat-

ing only in presence of appropriate fungus; seed

coat membranous, formed mainly by the outer

layer of the outer integument and generally con-

sists of one layer of cells; the inner integument

may persist as a thick layer; endosperm extremely

reduced and mostly lacking; embryo usually not

differentiated into distinct organs, and the cotyle-

don more or less develops only in a few species;

on germination the embryo forms a tubercle (pro-

tocorm) with basal rhizoids but without radicle

and usually without cotyledon. Produce ﬂ avone

C-glycosides and ﬂ avonols as major leaf constitu-

ents, n = 6–29 +.. . . . . . . . . . . . 5. orchidaceae.

3 Style and stamens free.

4 Capsules many-seeded. Herbs with more or

less tuberous rhizome or a corm often cov-

ered with membranous or ﬁ brous sheaths of

former leaves. Calcium oxalate raphides

generally present, silica bodies always

absent. Sieve-element plastids of form-P2c.

Leaves basal, mostly tristichous, linear to

lanceolate, prominently parallel-veined,

with mucilage canals, often clothed with

long, uniseriate hairs, but branched, multi-

cellular hairs also present. Stomata para-

cytic or rarely tetracytic. Flowers in axillary

racemes, spicate, corymbose or umbellate,

sometimes solitary, bisexual (rarely unisex-

ual in Curculigo), actinomorphic, 3-mer-

ous. Perianth segments six in two cycles

(rarely four in two cycles), spreading,

almost of equal size and similar in colour,

free or less often connate into a sometimes

long and narrow tube on top of the ovary,

more or less persistent. Stamens six in two

cycles or rarely (Pauridia) only three and

then opposite the inner segments; ﬁ laments

subulate to ﬁ liform, short to almost lacking;

arising from the base of perianth segments

or from the mouth of the perianth tube

anthers elongate, tetrasporangiate, basiﬁ xed

to dorsiﬁ xed, latrorse or introrse, opening

Superorder LILIANAE 647

longitudinally. Tapetum plasmodial (amoe-

boid). Microsporogenesis successive.

Pollen grains 2-celled, tectate-columellate,

1-colpate, ﬁ nely reticulate. stylodia connate

into short style that is apically divided into

three branches; ovary inferior, 3-locular to

1-locular (Empodium). Ovules often numer-

ous, anatropous, hemitropous, or campy-

lotropous, bitegmic, crassinucellate or

tenuinucellate, without parietal cell. Female

gametophyte of Polygonum-type or rarely

of Allium-type. Endosperm helobial or less

often (Pauridia and some species of Hypoxis

section Ianthe) nuclear; chalazal cell of

helobial endosperm somewhat haustorial.

Fruits either a capsule with loculicidal or cir-

cumscissile dehiscence, or sometimes more

or less ﬂ eshy and indehiscent or irregularly

dehiscent. Seeds small, brown or black, usu-

ally more or less globose, tuberculate to

spinulose, with a prominent raphe, often

with an elaiosome; seed coat is formed by

both integuments, but the most prominent

part is the outer epidermis of testa, the outer

cells of which develop characteristic protu-

berances; exotesta is characterized by prom-

inent, thick, black phytomelan crust; the

pigments mainly being stored in the cell

walls of the palisadlike exotesta, phloba-

phenes present also in tegmen; endosperm

composed of thin-walled, isodiametric cells

storing aleurone and fatty oils; embryo cylin-

drical, straight, usually more or less under-

developed, occupies two thirds or one half of

the length of seed. Alkaloids and steroidal

saponins absent but chelidonic acid present,

n = 6–9, 11. . . . . . . . . . . .4. hypoxidaceae.

4 Capsules 1-seeded. Tough herbs with rela-

tively short vertical rhizome with upper part

bearing several leaves and below them a coat

of densely set, ﬁ brous old leaf sheaths. Stem

with white, plumose hairs above, increas-

ingly dense on distal parts and on the inﬂ o-

rescences, this appearing white and woolly.

Leaves linear, stiff, basal, sheathing, gla-

brous. Stomata paracytic. Flowers in much

branched, dense, bracteate corymbose pani-

cle with rhipidial lateral units. Perianth

densely plumose, pubescent abaxially, gla-

brous adaxially, its segments connate into a

cylindrical- infundibular tube. Stamens all

fertile, inserted at the mouth of the tube;

anthers dorsiﬁ xed, versatile, introrse, basally

sagittate. Pollen grains 1-colpate, tectate-

columellate. Style ﬁ liform, apically 3-lobed;

ovary inferior or slightly semi-inferior, with

two ovules per locule, with septal nectaries.

The fertile locule dehiscing loculicidally.

Seeds black, shiny; seed coat mainly of outer

integument, with thick-walled, palisadlike

epidermal cells encrusted with phytomelan,

about six layers of testa below the epidermis

and strongly compressed two cell layers of

inner integument; endosperm copious, non-

starchy; embryo slightly curved, about as

long as endosperm. n = 9. 3. lanariaceae.

1. BLANDFORDIACEAE

R. Dahlgren et H. Clifford 1985. 1/4. Eastern Australia.

Blandfordia.

Probably the most archaic member of the order.

2. ASTELIACEAE

Dumortier 1829. 5/55. Mascarenes, New Guinea,

Australia, Tasmania, New Zealand, New Caledonia,

Fiji, Samoa, and Polynesia to Hawaii and Chile.

Astelia, Collospermum, Cordyline (including

Cohnia), Neoastelia, Milligania.

Shows some afﬁ nities with Blandfordiaceae and

Hypoxidaceae (Bayer et al. 1998; Rudall et al. 1998).

3. LANARIACEAE

H. Huber ex R. Dahlgren 1988. 1/1. Cape Province of

South Africa.

Lanaria.

Possible related to Asteliaceae and Hypoxidaceae

(Chase et al. 1995).

4. HYPOXIDACEAE

R. Brown 1814. 9/100–120. Africa (mainly South

Africa), Mascarenes, Seychelles, tropical and subtropical

648 Subclass II. LILIIDAE

regions of Asia (in north to eastern Himalayas, south-

western China, and Japan). New Guinea, Australia,

Tasmania, New Zealand, America from Atlantic part

of the USA to West Indies and Uruguay.

Hypoxis, Rhodohypoxis, Spiloxene, Pauridia,

Curculigo, Molineria, Hypoxidia, Empodium, Saniella.

Hypoxidaceae have probably some afﬁ nity with the

Asteliaceae (Rudall et al. 1997; Nordal 1998). Although

Hypoxidaceae and Asteliaceae differ from each other

in several respects, including successive microsporo-

genesis, inferior ovary and pollen sculpturing, there

are some unusual anatomical characters linking the

two families, notably branched trichomes and muci-

lage canals in the leaves (Rudall et al. 1998).

5. ORCHIDACEAE

A.L. de Jussieu 1789 (including Apostasiaceae Lindley

1833; Cypripediaceae Lindley 1833; Limodoraceae

Horaninow 1847; Neottiaceae Horaninow 1834;

Neuwiediaceae R. Dahlgren ex Reveal et Hoogland

1991; Vanillaceae Lindley 1835). 750–800/20000–

25000. Subcosmopolitan, chieﬂ y in tropical and sub-

tropical Asia, where representatives of all subfamilies

and tribes occur, and tropical America.

5.1 APOSTASIOIDEAE

Terrestrial, sometimes (Apostasia) rhizomatous herbs.

Vessels predominantly with simple perforations. Stems

woody at the base. Leaves spirally arranged, entire,

plicate. Stomata paracytic or tetracytic. Flowers

slightly zygomorphic, resupinate in Neuwiedia.

Labellum shallow. Stamens three (Neuwiedia), repre-

senting the median (adaxial) of the outer cycle and the

lateral ones of the inner cycle, or two (Apostasia), rep-

resenting lateral ones (the median stamen staminodial

or missing); ﬁ laments only partly fused with the style

in the gynostemium. Pollen grains free, 1-colpate with

an operculum, reticulate. Style with 2- or 3-lobed

stigma; stigma lobes equal and similar. Ovary 3- locular.

Fruits thin-walled or ﬂ eshy capsules, disintegrating or

(Neuwiedia veratrifolia) dehiscing loculicidally by

three valves. Seeds ovoid or elliptic, dark brown,

usually alveolate to reticulate, often sticky, sometimes

with long funicular appendage or with a prominent

appendage at both ends, n = 24. Southeastern Asia and

from the Himalayas and Ceylon to New Guinea and

northern Queensland. – Neuwiedia, Apostasia.

5.2 VANILLOIDEAE

Plant sympodial or monopodial. Vessels with scalari-

form perforation and the mostly opposite pitting of the

lateral walls. Stomata tetracytic. Staminodes two.

Anthers incumbent (by massive expansion of the api-

cal column/connective). Pollen grains in tetrads. Seeds

fusiform, crustose, n = 9, 10, 12, 14–16, 18. – Cleistes,

Isotria, Pogonia, Pogoniopsis, Duckeella, Erythrorchis,

Cyrtosia, Galeola, Pseudovanilla, Clematepistephium,

Dictyophyllaria, Epistephium, Eriaxis, Vanilla,

Lecanorchis.

5.3 CYPRIPEDIOIDEAE

Terrestrial or less often epiphytic herbs with ﬂ eshy or

ﬁ brous roots on short rhizomes. Vessels in roots or

rarely in stems, with simple or scalariform perfora-

tions. Leaves spirally arranged or distichous, thin and

plicate or ﬂ eshy and conduplicate. Stomata anomo-

cytic or paracytic. Flowers strongly zygomorphic,

resupinate. Two lateral sepals connate. Lateral petals

often considerably longer and narrower than the oth-

ers, median petal (labellum) deeply saccate, character-

istically slipper-shaped. Two lateral stamens of the

inner cycle always present, and the median stamen of

the outer cycle is modiﬁ ed into a characteristic shield-

like staminodium. Filaments largely united with style,

forming a thick, inﬂ exed gynostemium. Anthers sub-

globose and latrorse. Pollen grains in tetrads, usually

free, 1-colpate, ulcerate or porate, with various orna-

mentation; pollen more or less viscid, but pollinia

occur only in Phragmipedium longifolium and

Selenipedium chica. Style thick, the stigma large and

domelike. Ovary 3-locular or 1-locular. Fruits cap-

sules. Seeds subspheric with hard testa (Selenipedium)

or minute with membranous testa, n = 9 or more.

Eurasia, America. – Cypripedium, Selenipedium,

Mexipedium, Paphiopedilum, Phragmipedium.

5.4 NEOTTIOIDEAE

Mostly terrestrial herbs without pseudobulbs, often

facultative geophytes. Fertile stamen one or rarely

(Diplandorchis) two, the median one of both cycles.

Anther movably attached to gynostemium through a

connective, commonly persistent and soon withering,

the thecae usually closely approximate, parallel.

Pollinia granular, sectile or occasionally mealy,

2. – epipactideae: Cephalanthera, Sinorchis, Tangt-

sima, Epipactis, Limodorum, Epipogium, etc; neot-

tieae: Listera, Neottia, Diplandrorchis, Goodyera,

Superorder LILIANAE 649

Ludisia, Anoectochilus, Zeuxine, Spiranthes, Cranichis,

Ponthieva, etc.

5.5 EPIDENDROIDEAE. (including Vandoideae)

Mostly facultative epiphytes. Fertile stamen always 1.

Anther movably attached to gynostemium through a

connective, commonly deciduous. Pollinia waxy or

cartilaginous, rarely mealy, 2, 4, 6, or 8. – arethu-

seae: Arethusa, Sobralia, Elleanthus, Phaius,

Calanthe, Bletia, etc; gastroideae: Nervilia,

Gastrodia, etc.; epipogieae: Epipogium, Stereosandra;

coelogyneae: Coelogyne, Pleione, etc; malaxideae:

Liparis, Malaxis, Hammarbya, Oberoma, etc.; calyp-

soeae: Calypso, Yoania; epidendreae: Epidendrum,

Encyclia, Laelia, Cattleya, Eria, Pleurothallis, Stelis,

Masdevallia, Restrepia, Dracula, Dendrobium,

Flickingeria, Bulbophyllum, etc.; polystachyeae:

Polystachya, etc.; cymbidieae: Eulophia, Ansellia,

Cymbidium, Grammatophyllum, Catasetum,

Stanhopea, Gongora, Oncidium, Odontoglossum,

Miltonia, Brassia, etc.; maxillarieae: Corallorrhiza,

Zygopetalum, Lycaste, Maxillaria, etc.; vandeae:

Phalaenopsis, Aerides, Vanda, Arachnis, Renanthera,

Cleisostoma, Aerangis, Taeniophyllum, Microcoelia,

Angraecum, etc.

5.6 ORCHIDOIDEAE

Usually terrestrial herbs with spirally arranged leaves.

Vessels in roots, with scalariform perforation.

Commonly with storage organ, the tuberoid, which is a

root structure around a core of stem tissue with an api-

cal bud that will grow into a new shoot in the next sea-

son, with one of the axillary buds forming a new

tuberoid. Fertile stamen one. Anther ﬁ rmly adnate to

gynostemium, not deciduous or withering. Thecae

often widely spaced, parallel, or converging. Pollinia

rather soft, with caudicles, the caudicles directed

toward base of anther. – orchideae: Orchis,

Dactylorhiza, Ophrys, Anacamptis, Platanthera,

Gymnadenia, Habenaria, Cynorkis, Peristylus, etc;

cranichideae: Cyanicula, Aspidogyne, Moerenhoutia,

Zeuxine, Beloglottis, Spiranthes, Cranichis, Manniella,

etc.; diurideae: Acianthus, Corybas, Cyrtostylis,

Bipinnula, Chloraea, Gavilea, Pterostylis,

Stigmatodactylus, Adenochilus, Caladenia,

Elythranthera, Eriochilus, Glossodia, Leptoceras,

Ptaecoxanthus, Coilochilus, Diuris, Orthoceras,

Caleana, Chiloglottis, Drakaea, Leporella, Lyper-

anthus, Megastylis, Paracaleana, Pyrorchis, Rumicola,

Spiculaea, Genoplesium, Microtis, Thelymitra,

Rhizanthella, Prasophyllum, Calochilus; diseae:

Disa, Satyrium, Disperis, etc.

A rather taxonomically isolated group, which is

characterized by a number of unique characters, includ-

ing miniaturization of the ovules and microspermy,

silica bodies, and mesophyll idioblasts with helical

thickening. They show connections with the

Hypoxidaceae (Rolfe 1909, 1910; Hallier 1912; Gobi

1916; Hutchinson 1934, 1959, 1973; Garay 1960; Rao

1969; Takhtajan 1980, 1987, 1997; Goldblatt 1995),

especially with the genera Hypoxis and Curculigo,

there are indeed many similarities between Orchidaceae

and Hypoxidaceae, and Curculigo orchioides, for

example, strongly resemble the Orchidaceae. Also,

chromosomes of the Hypoxidaceae resemble those of

the Orchidaceae (Sharma 1969). Evidence from rbcL

sequences also indicates that orchids are most closely

related to Hypoxidaceae (Cameron et al. 1994). The

connecting link between Hypoxidaceae and

Orchidaceae is the most archaic subfamily the

Apostasioideae. This intermediate position of

apostasiads was already mentioned by Lindley (1833)

in his The Vegetable Kingdom (“The order seems as if

connecting Orchids with Hypoxids”). However,

Orchidaceae could not be derived directly from the

Hypoxidaceae.

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Order 13. IRIDALES

Mainly perennial herbs, generally with sympodial

rhizomes or corms, less often with bulbs, occasion-

ally shrubs with woody caudex and anomalous sec-

ondary growth, very rarely annuals (Sisyrhinchium

spp.). Roots often mycorrhizal, without root hairs.

Vessels usually only in roots (except Sisyrhinchium),

with scalariform or simple perforations. Raphides

lacking, but styloid calcium oxalate crystals present

in vascular bundles and other tissues (absent only in

Sisyrhinchium and closely related genera). Leaves

basal or basal and cauline, generally distichous, often

ensiform, linear or sometimes lanceolate or ﬁ liform,

rarely (Doryanthaceae) very large and long (to 3 m

long), rarely (Geosiris) scalelike, sheathing and

equitant at the base, parallel-veined. Stomata anomo-

cytic or paracytic. Flowers in terminal, cymose inﬂ o-

rescences or sometimes solitary, usually bracteate,

mostly bisexual, actinomorphic or often more or less

zygomorphic. Perianth of six petaloid segments in

two cycles, all alike or the two cycles or the members

of a single cycle more or less conspicuously different

in shape and colour, often all connate below into a

tube; occasionally the inner cycle reduced or lacking.

Nectaries perigonal (Iridoideae), septal (Ixioideae

and Nivenioideae) or sometimes lacking (e.g.

Isophysis, most Aristea). Stamens six in two cycles or

three (two in Diplarrhena); ﬁ laments free or some-

times connate basally or entirely; anthers basi ﬁ xed to

subbasiﬁ xed, or centriﬁ xed, tetrasporangiate, extrorse

to latrorse, usually opening longitudinally, rarely

opening by apical pores. Tapetum secretory.

Microsporogenesis mostly simultaneous. Pollen

grains mostly 2-celled, tectate-columellate, usually

654 Subclass II. LILIIDAE

1-colpate, rarely 2- or 3-colpate, spiraperturate, or

inaperturate, usually reticulate. Gynoecium of three

united carpels; style terminal, subulate to ﬁ liform,

usually 3-branched above, the branches often again

divided, sometimes expanded and strongly petaloid

with the stigma on the outer side of the branch rather

than at the top; ovary inferior or very rarely (Isophysis)

superior, 3-locular or very rarely (Hermodactylis)

1-locular with parietal placentation, usually with

numerous ovules per locule, rarely with few or only

one ovule. Ovules anatropous or campylotropous,

bitegmic, crassinucellate, with parietal tissue and

sometimes (Iris pseudacorus) with endothelium and

in some genera with funicular obturator. Female

gametophyte of Polygonum-type. Endosperm helo-

bial (Doryanthes, Ixiolirion, Lanaria, Odontostomum,

Geosiris and Isophysis) or nuclear. Fruits loculicidal

capsules, rarely indehiscent. Seeds sometimes aril-

late; embryo rather small, cylindrical, mostly straight;

both testa and tegmen are usually present; the outer

epidermis is mostly pigmented with phlobaphene;

endosperm hard, with thickened, pitted cell walls that

contain reserves of hemicellulose; cells also contain

aleurone and lipids but only rarely starch; embryo

straight and fusiform and is usually 1/3–2/3 as long

as the endosperm. The cotyledon of the seedling is

coleoptilelike. Produces ﬂ avone C-glycosides and

ﬂ avonols as major leaf constituents.

The distinctness of the Iridales is well supported by

several characters, one of which are unique – calcium

oxalate crystals in the form of styloids (Goldblatt et al.

1984). Probably related to the Melanthiaceae s.

l. (Takhtajan 1959, 1966, 1969, 1980, 1997; Goldblatt

1990), and to the Campynemataceae. Embryologically

the closest to the Melanthiales is Geosiris (Goldblatt

1990).

Key to Families

1 Microsporogenesis successive. Endosperm helobial.

2 Herbs with short vertical rootsock and large leaves.

Leaves in dense basal rosette, from 1.3 to 3 m long

and 20 cm wide, tough, swordlike, those on the

stem much reduced, each with characteristic brown

tubular tip. Mesophyll containing calcium oxalate

crystals. Stomata paracytic. Inﬂ orescences tall (up

to 5 m long) terminal scapes, oblong thyrses or

global compound racemes of numerous ﬂ owers,

which may be substituted by bulbils. Flowers large

(5–10 cm long), bisexual, actinomorphic or slightly

zygomorphic. Perianth red, segments nearly free

or basally connate into very short tube, long and

falcate. Filaments adnate to perianth segments for

about one-half of their length; anthers elongate,

basiﬁ xed (the apex of ﬁ lament inserted into a pit at

base of connective), opening longitudinally.

Microsporogenes successive. Pollen grains 1-col-

pate. Ovary inferior, 3-locular, with 40–50 anatro-

pous ovules per locule; styles two long, tapering,

with triangular stigma. Septal nectaries present.

Endosperm helobial. Fruits large, turbinate, loculi-

cidal capsules. Seeds light brown, more or less ﬂ at-

tened, elliptic to rhomboid, with one-sided, broad,

papery wing, nonligniﬁ ed exotestal cell walls of

Doryanthes palmeri impregnated with phloba-

phene; embryo linear, from one-third to almost as

long as the seed and with broad cotyledon. Testa

without phytomelan. Proanthocyanidins and ste-

roid saponines present. n = 17, 18, 22, 24, with

clearly detectable differences in chromosome

sizes.. . . . . . . . . . . . . . . . . . . .5. doryanthaceae.

2 Small, achlorophyllous, mycotrophic herbs with

mycorrhizal roots. Vessels in roots, with scalari-

form perforations. Leaves small, scalelike, alter-

nate, membranous, simple, entire. Stomata

anomocytic. Flowers in terminal cymes, bracte-

ate, small, sessile, actinomorphic. Perianth seg-

ments six in two whorled (the outer imbricate,

the inner contorted), petaloid, blue; perigone

tube short. Stamens three, opposite the outer

perianth segments, very short; anthers basiﬁ xed,

extrorse, opening longitudinally. Pollen grains

1-colpate. Septal nectaries absent. Ovary infe-

rior, 3-locular; style one, trilobed, stigmas three;

ovules anatropous, placentation axile. Fruits cap-

sules or capsular-indehiscent, capsules triangu-

lar-obconical. Seeds minute, testa without

phytomelan.. . . . . . . . . . . . . . 6. geosiridaceae.

1 Microsporogenesis simultaneous. Endosperm

mostly nuclear.

3 Stamens mostly six, all fertile or 2–3 stamino-

dial, attached to segments.

4 Ovules without chalazosperm.

5 Anthers opening longitudinally. Glabrous

herbs with leafy stem and bulb-shaped tuni-

cated corm that develops lateral corms at

some distance from its base. Vessels in roots

with scalariform perforations. Leaves linear,

not stiff, cylindrical-subulate at the apex.