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Family Liliaceae

Liliaceae Juss.

Including Fritillariaceae Salisb., Medeolaceae (S. Watson) Takhtajan, Tulipaceae Batsch ex BorkkhausenExcluding Alliaceae J.G. Agard, Alstroemeriaceae Dum, Amaryllidaceae Jaume St.-Hil, Anthericaceae J.G. Agard, Aphyllanthaceae G.T. Burnet, Asparagaceae Juss, Asphodelaceae Juss, Asteliaceae Dum, Blandfordiaceae Dahlgren and Cliffor, Calochortaceae Dum, Colchicaceae DC, Convallariaceae Horan, Eriospermaceae Endl, Hemerocallidaceae R. Br, Herreriaceae, Hostaceae B. Mathe, Hyacinthaceae J.G. Agard, Hypoxidaceae R. Br, Melanthiaceae Batsc, Ruscaceae Spreng, Tecophilaeaceae Leybol, Trilliaceae Lindl.

Habit and leaf form. Herbs. ‘Normal’ plants, or switch-plants; occasionally phyllodineous. Leaves well developed (usually), or much reduced (infrequently). Plants non-succulent. Perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Young stems not breaking easily at the nodes. Bulbaceous. Helophytic to xerophytic. Leaves alternate (nearly always), or opposite, or whorled; when alternate, spiral (always?); flat, or folded, or rolled, or terete; sessile, or petiolate; sheathing, or non-sheathing. Leaf sheaths with free margins. Leaves simple; epulvinate. Lamina entire; linear to lanceolate, or ovate; parallel-veined; without cross-venules. Leaves eligulate.

General anatomy. Accumulated starch other than exclusively ‘pteridophyte type’.

Leaf anatomy. Stomata present; anomocytic.

Lamina dorsiventral, or centric. The mesophyll without calcium oxalate crystals (according to Goldblatt 1995). Minor leaf veins without phloem transfer cells (2 genera). Vessels absent.

Stem anatomy. Secondary thickening absent. Xylem without vessels. Sieve-tube plastids P-type; type II.

Root anatomy. Roots with velamen (e.g. in Lilium), or without velamen. Root xylem with vessels; vessel end-walls scalariform.

Reproductive type, pollination. Fertile flowers hermaphrodite. Unisexual flowers absent. Plants hermaphrodite. Floral nectaries present. Nectar secretion from the perianth (via nectaries at the tepal bases). Pollination entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; in racemes, or in umbels, or in panicles. The ultimate inflorescence unit racemose. Inflorescences scapiflorous, or not scapiflorous; terminal, or axillary; espatheate. Flowers bracteate, or ebracteate; ebracteolate (usually), or bracteolate (occasionally); small to large; regular (nearly always), or somewhat irregular; sometimes somewhat zygomorphic; 3 merous; cyclic; pentacyclic. Perigone tube absent.

Perianth of ‘tepals’ (usually), or with distinct calyx and corolla; 6; free; 2 whorled; isomerous; petaloid, or sepaloid and petaloid; without spots, or spotted (commonly); similar in the two whorls, or different in the two whorls (the outer segments often smaller and less showy); colour variable.


Androecium 6. Androecial members free of the perianth; all equal; free of one another; 2 whorled (3+3). Androecium exclusively of fertile stamens. Stamens 6; diplostemonous; alterniperianthial. Anthers (pseudo) basifixed (often, the filament tip enclosed by a tubular outgrowth from the back of the connective), or dorsifixed; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. The endothecial thickenings spiral. Microsporogenesis successive. Anther wall initially with more than one middle layer; of the ‘monocot’ type. Tapetum glandular. Pollen grains aperturate; 1 aperturate; usually sulcate (in Tulipa occasionally operculate, or with an irregular aperture); 2-celled.

Gynoecium 3 carpelled. Carpels isomerous with the perianth. The pistil 3 celled. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 3 locular. Styles 1; attenuate from the ovary, or from a depression at the top of the ovary; apical. Stigmas 1, or 3; dorsal to the carpels; wet type, or dry type; papillate; Group II type, Group III type, and Group IV type. Placentation axile. Ovules 5–50 per locule (usually ‘many’); arillate, or non-arillate; anatropous; bitegmic (without a parietal cell); tenuinucellate. Outer integument not contributing to the micropyle. Embryo-sac development Fritillaria-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Endosperm formation helobial. Embryogeny onagrad.

Fruit non-fleshy; dehiscent; a capsule. Capsules loculicidal. Seeds endospermic. Endosperm oily. Seeds winged (e.g., Lilium), or wingless. Seeds without starch. Embryo well differentiated (small). Cotyledons 1 (not coleoptile-like). Embryo achlorophyllous (two species, representing Fritillaria and Tulipa). Testa without phytomelan; brown or pallid.


Seedling. Hypocotyl internode present, or absent. Seedling collar not conspicuous. Cotyledon hyperphyll elongated (mostly), or compact (sometimes in Lilium); assimilatory (mostly), or non-assimilatory (sometimes in Lilium); more or less circular in t.s. (mostly), or dorsiventrally flattened (Lilium). Coleoptile absent. Seedling cataphylls absent. First leaf dorsiventral. Primary root ephemeral.

Physiology, biochemistry. Not cyanogenic. Alkaloids present, or absent. Proanthocyanidins absent. Flavonols present (mostly), or absent; quercetin, or kaempferol and quercetin. Ellagic acid absent. Saponins/sapogenins present, or absent. C3. C3 physiology recorded directly in Zigadenus. Anatomy non-C4 type (Zigadenus).

Geography, cytology. Holarctic, Paleotropical, and Neotropical. Northern hemisphere, centred on southwest and Himalayan Asia to China. X = 12.

Taxonomy. Subclass Monocotyledonae. Dahlgren et al. Superorder Liliiflorae; Liliales. APG 3 core angiosperms; Superorder Lilianae; non-commelinid Monocot; Order Liliales.

Species about 420. Genera about 10; Cardiocrinum, Erythronium, Fritillaria, Gagea, Lilium, Lloydia, Medeola(?), Nomocharis, Notholirion(?), Tulipa.

Illustrations.
• Technical details: Fritillaria imperialis.
• Erythronium grandiflorum: Bot. Reg. 1786, 1836.
• Fritillaria meleagris (B. Ent.).
• Fritillaria spp. (Chittenden).
• Galtonia, Lilium, Nomocharis.
• Lilium martagon: Eng. Bot. 1518 (1869).
• Lilium x testaceum (L. candidum x L. chalcedonicum): Bot. Reg. 29, 11 (1843).
• Lloydia serotina: Eng. Bot. 1521 (1869).
• Maianthemum bifolium: as Smilacina bifolia, Eng. Bot. 1510 (1869).
• Tulipa “gesneriana”: Bot. Reg. XXIV, 46 (1838).
• Tulipa maleolens: Bot. Reg. 1839, 66.
• Tulipa sylvestris (B. Ent.).
• Tulipa sylvestris: Eng. Bot. 1520 (1869).


Quotations

To gild refinèd gold, to paint the lily
. . . . . . .
Is wasteful, and ridiculous excess
(‘King John’, iv., 2)

O sweetest, fairest lily!
My brother wears thee not one half so well
As when thou grewest thyself
(‘Cymbeline’, iv., 2)

I like the chaliced lilies,
The heavy Eastern lilies,
The gorgeous tiger-lilies,
That in our garden grow!
(T.B. Aldrich, ‘Tiger-Lilies’)

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