| | The Families of Angiosperms |
Including Fritillariaceae Salisb., Medeolaceae (S. Watson) Takhtajan, Tulipaceae Batsch ex Borkkhausen; excluding 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. Mathew, 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 green and photosynthesizing. Perennial; with a basal aggregation of leaves, or without conspicuous aggregations of leaves; bulbaceous, or cormous, or rhizomatous, or tuberous. Helophytic to xerophytic. Leaves alternate (nearly always), or opposite, or alternate and 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.
Leaf anatomy. The leaf lamina dorsiventral, or centric. Stomata present; anomocytic. The mesophyll without crystals (according to Goldblatt 1995). Foliar vessels absent. Minor leaf veins without phloem transfer cells (2 genera).
Axial (stem, wood) anatomy. Secondary thickening absent. The axial xylem without vessels.
Root anatomy. Roots with velamen (e.g. in Lilium), or without velamen. Root xylem with vessels; vessel end-walls scalariform.
Reproductive type, pollination. Unisexual flowers present, or absent. Plants hermaphrodite, or andromonoecious. 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’; when aggregated, in racemes, or in umbels, or in panicles. The ultimate inflorescence units racemose. Inflorescences scapiflorous, or not scapiflorous; terminal, or axillary. 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 3 (Scoliopus), or 6. Androecial members free of the perianth; free of the gynoecium; all equal; free of one another; 2 whorled (3+3). Androecium exclusively of fertile stamens. Stamens 6 (usually), or 3 (Scoliopus); reduced in number relative to the adjacent perianth, or diplostemonous (usually); 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; extrorse (Scoliopus), or 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 (usually), or 1 celled (but 3-lobed, Scoliopus). Gynoecium syncarpous; synstylovarious (Scoliopus), or eu-syncarpous (usually); superior. Ovary 3 locular. Gynoecium stylate. Styles 1; attenuate from the ovary, or from a depression at the top of the ovary; apical. Stigmas 1, or 3; wet type, or dry type; papillate; Group II type, Group III type, and Group IV type. Placentation in Scoliopus, parietal; usually 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 not conspicuously hairy; 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, phytochemistry. C3. C3 physiology recorded directly in Zigadenus. Anatomy non-C4 type (Zigadenus). Accumulated starch other than exclusively ‘pteridophyte type’. Not cyanogenic. Alkaloids present, or absent. Saponins/sapogenins present, or absent. Proanthocyanidins absent. Flavonols present (mostly), or absent; quercetin, or kaempferol and quercetin. Ellagic acid absent. Sieve-tube plastids P-type; type II.
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 III core angiosperms; Superorder Lilianae; non-commelinid Monocot. APG IV Order Liliales.
Species about 420. Genera about 10; Cardiocrinum, Clintonia, Erythronium, Fritillaria, Gagea, Lilium, Lloydia, Medeola, Nomocharis, Notholirion(?), Prosartes, Scoliopus(?), Speirantha(?), Streptopus, Tulipa, Tricyrtis.
General remarks. Our compiled data strongly suggest that current taxonomic assignment of Scoliopus to Liliaceae sensu stricto (e.g., Christenhusz et al., 2017) is very unsatisfactory. This North American genus calls for direct examination of the two species to assess and perhaps re-interpret the peculiar floral morphology. Descriptions in modern floras (three fertile stamens only, attached to the ‘sepals’ and alternating with narrow linear ‘petals’, and a unilocular G3 with parietal placentation) conform with J.D. Hooker's treatment in Bot. Mag. 123 (1897), from which our illustraton is adapted.
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’)
Illustrations. • Le Maout and Decaisne: Fritillaria imperialis. • Erythronium californicum (as E. giganteum): Bot. Mag. 94 (1868). • Erythronium hendersoni: Bot. Mag. 114 (1888). • Erythronium multiscapidium (as E. hartwegi): Bot. Mag. 124 (1898). • Erythronium grandiflorum: Bot. Reg. 1786, 1836. • Fritillaria aurea: Bot. Mag. 120 (1894). • Fritillaria bucharica: Bot. Mag. 115 (1889). • Fritillaria caucasica (as F. tulipifolia): Bot. Mag. 98 (1872). • Fritillaria latifolia (as F. nobilis): Bot. Mag. 122 (1896). • Fritillaria raddeana (as F. askabadensis): Bot. Mag. 128 (1902). • Fritillaria meleagris (B. Ent.). • Fritillaria spp. (Chittenden). • Fritillaria pluriflora: Bot. Mag. 124 (1898). • Fritillaria pontica: Bot. Mag. 146 (1920). • Fritillaria recurva: Bot. Mag. 102 (1876). • Galtonia, Lilium, Nomocharis. • Lilium bakerianum var. bakerianum (as L. lowii): Bot. Mag. 118 (1892). • Lilium duchartrei (as L. farreri): Bot. Mag. 146 (1920). • Lilium henryi: Bot. Mag. 117 (1891). • Lilium japonicum var. japonicum (as Lilium krameri): Bot. Mag. 99 (1873). • Lilium martagon: Eng. Bot. 1518 (1869). • Lilium oxypetalum (as Fritillaria): Bot. Mag. 79 (1853). • Lilium rubellum: Bot. Mag. 125 (1899). • Lilium x testaceum (L. candidum x L. chalcedonicum): Bot. Reg. 29, 11 (1843). • Lloydia serotina: Eng. Bot. 1521 (1869). • Lloydia tibetica: Hook. Ic. Pl. 23 (1894). • Maianthemum bifolium: as Smilacina bifolia, Eng. Bot. 1510 (1869). • Notholirion thomsonianum (as Lilium roseum): Bot. Mag. 79 (1853). • Prosartes smithii (as Disporum): Abrams, Ill. Flora of the Pacific States 1 (1923). • Scoliopus bigelovii: Bot. Mag. 123 (1897). • Speirantha gardenii (as Albuca): Bot. Mag. 81 (1855). • Tricyrtis hirta: Bot. Mag. 89 (1863). • Tricyrtis macropoda: Bot. Mag. 107 (1881). • Tricyrtis stolonifera: Bot. Mag. 140 (1914). • Le Maout and Decaisne: Tricyrtis. • Tulipa agenensis (as T. maleolens): Bot. Reg. 1839, 66. • Tulipa gesneriana: Bot. Reg. XXIV, 46 (1838). • Tulipa orphanidea (as T. haeri): Bot. Mag. 102 (1876). • Tulipa humilis var. pulchella (as T. pulchella): Bot. Mag. 103 (1877). • Tulipa orphanidea: Bot. Mag. 103 (1877). • Tulipa praestans: Bot. Mag. 129 (1903). • Tulipa sylvestris (B. Ent.). • Tulipa sylvestris: Eng. Bot. 1520 (1869). • Tulipa tetraphylla (as T. kesselringii): Bot. Mag. 110 (1884).
We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, genera included in each family, and classifications (Dahlgren; Dahlgren, Clifford, and Yeo; Cronquist; APG). See also Guidelines for using data taken from Web publications.
Cite this publication as: ‘Watson, L., and Dallwitz, M.J. 1992 onwards. The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 4th May 2024. delta-intkey.com’.
