Family Amaryllidaceae

Amaryllidaceae Jaume St.-Hil.

~ Liliaceae

Including Brunsvigiaceae Horan., Cepaceae Salisb.(?), Cyrtanthaceae Salisb., Galanthaceae Salisb., Gethyllidaceae Rafin., Operantheae (Operanthaceae) Salisb., Leucojaceae Batsch & Borkhausen, Strumariaceae Salisb., Zephyranthaceae Salisb.

Excluding Doryanthaceae, Hypoxidaceae

Habit and leaf form. Herbs (without allylic sulphides). Perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves; bulbaceous (mostly), or rhizomatous (in that a few have structures transitional between rhizomes and bulbs). Mesophytic. Leaves mostly deciduous; alternate; spiral (not uncommonly, e.g. Crinum), or distichous (mostly?); ‘herbaceous’; sessile, or petiolate (or almost so); sheathing; without marked odour (in particular, not onion-scented); simple. Lamina entire; linear, or lanceolate, or oblong, or ovate, or orbicular; parallel-veined; without cross-venules. Lamina margins entire. Leaves with a persistent basal meristem, and basipetal development.


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

Leaf anatomy. Stomata present; anomocytic.

Lamina dorsiventral. The mesophyll containing mucilage cells (with raphides); containing calcium oxalate crystals. The mesophyll crystals raphides. Minor leaf veins without phloem transfer cells (Amaryllis, Zephyranthes). Vessels absent.

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

Root anatomy. 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 (from the inner tepals, in Galanthieae), or from the gynoecium (from septal nectaries).

Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’; in cymes (variously condensed), or in umbels, or in heads. The ultimate inflorescence unit cymose. Inflorescences scapiflorous; cymose, but often umbels or heads by condensation; with involucral bracts (mostly with two(–8) spathelike, free or connate scales), or without involucral bracts; spatheate. Flowers regular to somewhat irregular to very irregular; when irregular, somewhat zygomorphic; 3 merous; cyclic; tetracyclic, or pentacyclic. Perigone tube present (short to long), or absent.

Perianth of ‘tepals’; 6; free to joined; 2 whorled (3+3, but often with a conspicuous ‘corona’, like an extra, inner whorl); isomerous; petaloid; similar in the two whorls; green, or green to white, or white, or cream, or yellow, or red, or pink, or purple, or brown (in various combinations, but not blue).

Androecium (3–)6(–18) (nearly always 3+3). Androecial members free of the perianth, or adnate (to the tube); free of the gynoecium; free of one another, or coherent; when joined 1 adelphous; nearly always 2 whorled (3+3). Androecium exclusively of fertile stamens (at least, reduction to staminodes not mentioned by Dahlgren et al. 1985). Stamens 3 (in Zephyra), or 6 (usually), or 9–18 (Gethyllis); isomerous with the perianth (rarely), or diplostemonous; (nearly always?) alterniperianthial; filantherous (the filaments sometimes appendaged alongside the anthers). Filaments appendiculate (the connate filaments sometimes expanded to form a staminal corona), or not appendiculate. Anthers dorsifixed (epipeltate), or basifixed (rarely); versatile (usually), or non-versatile; dehiscing via pores, or dehiscing via longitudinal slits; introrse (usually), or latrorse (e.g. Crinum); tetrasporangiate. Endothecium developing fibrous thickenings. The endothecial thickenings spiral. Microsporogenesis successive. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Tapetum glandular (usually), or amoeboid. Pollen grains aperturate; 1(–2) aperturate; sulcate (usually), or sulculate (Amaryllideae); 2-celled.

Gynoecium 3 carpelled. Carpels isomerous with the perianth. The pistil usually more or less 3 celled, or 1 celled (e.g., in Calostemma, and in other forms the dissepiments of an ostensibly trilocular ovary are merely contiguous). Gynoecium syncarpous; eu-syncarpous; inferior, or partly inferior (rarely). Ovary 3 locular (usually), or 1 locular (rarely, though not uncommonly approaching this condition). Gynoecium stylate. Styles 1; apical. Stylar canal present. Stigmas 1, or 3; 1–3 lobed; capitate; dry type (mostly), or wet type (some); papillate; Group II type and Group III type. Placentation when unilocular, basal (viz., in Calostemma,), or parietal; when trilocular (i.e. usually), axile, or basal (rarely, or at least ostensibly so with the ovules solitary or paired and collateral). Ovules (1–)12–50 per locule (i.e. nearly always ‘several to many’); non-arillate; anatropous; without integuments (rarely), or unitegmic, or bitegmic (usually); crassinucellate, or pseudocrassinucellate. Embryo-sac development Polygonum-type (usually), or Allium-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral, or persistent (often). Synergids hooked (with filiform apparatus). Hypostase present (Zephyranthes), or absent. Endosperm formation nuclear, or helobial.

Fruit fleshy, or non-fleshy; dehiscent, or indehiscent; a capsule, or a berry. Capsules loculicidal. Seeds endospermic. Endosperm oily. Seeds winged, or wingless. Seeds without starch. Cotyledons 1. Embryo achlorophyllous (5/5), or chlorophyllous (two species of Haemanthus); straight. Testa encrusted with phytomelan (mostly?), or without phytomelan (e.g., Amaryllis, Hymenocallis); black (mostly), or green, or red.

Seedling. Hypocotyl internode absent. Seedling collar not conspicuous. Cotyledon hyperphyll elongated, or compact; assimilatory, or non-assimilatory; when elongated, dorsiventrally flattened. Coleoptile absent. First leaf dorsiventral. Primary root ephemeral.

Physiology, biochemistry. Cyanogenic, or not cyanogenic. Alkaloids present (amaryllid type), or absent. Proanthocyanidins absent. Flavonols present, or absent; when present, quercetin, or kaempferol and quercetin. Ellagic acid absent. Saponins/sapogenins absent. Inulin not found (Gibbs 1974). C3. C3 physiology recorded directly in Zephyranthes. Anatomy non-C4 type (Zephyranthes).

Geography, cytology. Holarctic, Paleotropical, Neotropical, Cape, and Australian. Indian, Indo-Chinese, and Malesian. Temperate (a few), or sub-tropical and tropical (many). Widespread.

Taxonomy. Subclass Monocotyledonae. Superorder Liliiflorae; Asparagales. APG (1998) Monocot; non-commelinoid. APG 3 (2009) Order: Asparagales.

Species about 800. Genera about 60; Amaryllis, Ammocharis, Apodolirion, Bokkeveldia, Boophone, Bravoa, Brunsvigia, Caliphruria, Calostemma, Carpolyza, Chlidanthus, Choananthus, Clivia, Cooperia, Crinum, Cryptostephanus, Cybistetes, Cyrtanthus, Eucharis, Eucrosia, Eustephia, Galanthus, Gemmaria, Gethyllis, Griffinia, Habranthus, Haemanthus, Hannonia, Hessea, Hieronymiella, Hippeastrum, Hymenocallis, Ismene, Lapiedra, Leptochiton, Leucojum, Lycoris, Namaquanula, Narcissus, Nerine, Pamianthe, Pancratium, Paramongaia Phaedranassa, Phycella, Placea, Proiphys (Eurycles), Pucara, Pyrolirion, Rauhia, Rhodophiala, Scadoxus, Sprekelia, Stenomesson, Sternbergia, Strumaria, Tapeinanthus, Tedingea, Traubia, Ungernia, Urceolina, Vagaria, Vallota, Worsleya, Zephyranthes (Dahlgren et al. (1985) omit many genera).

Economic uses, etc. Many cultivated ornamentals.

Illustrations.
• Technical details: Leucojum.
• Technical details: Narcissus, Galanthus, Oporanthus.
• Narcissus, Galanthus, Leucojum (B. Ent. compilation).
• Amaryllis belladonna (~ Brunsvigia rosea?): as A. banksiana, Bot. Reg. 11, 1842.
• Calostemma luteum: Bot. Reg. xxvi, 19 (1840).
• Calostemma purpureum: as C. carneum, Bot. Reg. xxvi, 26 (1840).
• Crinum roseum: as C. variabile var. roseum, Bot. Reg. 1844, 9.
• Crinum: habit and LS flower (Thonner).
• Cyrtanthus carneus: Bot. Reg. 1462, 1831.
• Galanthus nivalis: Eng. Bot. 1507 (1869).
• Habranthus andersoni: Bot. Reg. 1345, 1830.
• Habranthus gracilifolius var. boothianus: Bot. Reg. 1967, 1837.
• Hippeastrum cybister: as Sprekelia, Bot. Reg. xxvi, 33 (1840).
• Hippeastrum pratense: as Habranthus pratensis, Bot. Reg. 35, 1842.
• Leucojum vernum: Bot. Mag. 2, 1788.
• Narcissus triandrus: Bot. Mag. 2, 1788.
• Narcissus poeticus: Eng. Bot. 1504 (1869).
• Narcissus pseudonarcissus: Eng. Bot. 1388 (1869).
• Narcissus pseudonarcissus ssp. major: Bot. Mag. 2, 1788.
• Assorted cultivated Narcissi: Bot. Reg. 29, 38 (1843).
• Pancratium - habit and technical details (Lindley).
• Phycella herbertiana: Bot. Reg. 1341, 1830.
• Placea ornata: Bot. Reg. 50, 1841.
• Pyrolirion aureum: Bot. Reg. 1724, 1835.
• Sprekelia formosissima: Bot. Mag. 2, 1788.
• Sprekelia formosissima: as S. glauca, Bot. Reg. 16, 1841.
• Stenomesson aurantiacum: as S. hartwegii, Bot. Reg. 1844, 42.
• Stenomesson flavum: Bot. Reg 29 (2), 1843.
• Urceolina peruviana: Bot. Reg. 1839, 68.

Quotations
. . . Daffodils
That come before the swallow dares, and take
The winds of March with beauty
(‘The Winter’s Tale’, iv., 3)

I wander’d lonely as a cloud
That floats on high o’er vales and hills,
When all at once I saw a crowd,
A host, of golden daffodils;
Beside the lake, beneath the trees,
Fluttering and dancing in the breeze.
(William Wordsworth, ‘Daffodils’)

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