Family Hyacinthaceae

Hyacinthaceae J.G. Agardh

~ Former Liliaceae-Scilloideae, cf. Asparagaceae-Scilloideae of APG III

Including Eucomidaceae Salisb., Lachenaliaceae Salisb., Scillaceae von Vest

Habit and leaf form. Herbs. ‘Normal’ plants. Perennial; with a basal aggregation of leaves; bulbaceous (usually), or rhizomatous (Chlorogalum, Schoenolirion). Mesophytic. Leaves alternate; spiral (always?); sessile; sheathing. Leaf sheaths with free margins. Leaves simple. Lamina entire; linear, or lanceolate, or ovate (rarely), or orbicular (rarely); parallel-veined; without cross-venules. Lamina margins entire.

Leaf anatomy. Stomata present; anomocytic.

The mesophyll containing mucilage cells (with raphides); usually containing calcium oxalate crystals. The mesophyll crystals raphides. Vessels absent.

Stem anatomy. Secondary thickening absent. Xylem without vessels.

Root anatomy. Root xylem with vessels; vessel end-walls scalariform, or scalariform and simple.

Reproductive type, pollination. Plants hermaphrodite, or polygamomonoecious. Floral nectaries present. Nectar secretion from the gynoecium (from septal nectaries).

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in racemes, in spikes, and in heads. The ultimate inflorescence unit racemose. Inflorescences scapiflorous; terminal; usually simple or branched racemes or spikes, rarely heads; espatheate. Flowers bracteate (at least in lower parts of the inflorescence); small, or medium-sized; regular (usually), or somewhat irregular, or regular and somewhat irregular, or regular and very irregular (occasionally in heads with large, very irregular ray florets and smaller, more or less regular disk florets - see illustration of Daubenya aurea); 3 merous; cyclic; pentacyclic. Perigone tube present (usually, campanulate, urceolate or tubular), or absent (e.g. Ornithogalum).

Perianth of ‘tepals’; 6; free, or joined; 2 whorled; isomerous; petaloid; similar in the two whorls, or different in the two whorls (but usually of the same texture); white, or yellow, or red, or violet, or blue, or brown, or black.

Androecium 6 (usually), or 3 (e.g. in Albuca, where the outer whorl may be reduced or absent). Androecial members free of the perianth, or adnate (to the tube); free of one another; 2 whorled (usually), or 1 whorled (sometimes, in Albuca). Androecium exclusively of fertile stamens (usually), or including staminodes (e.g., sometimes in Albuca). Staminodes when present, 3; when present, external to the fertile stamens. Stamens 6 (usually), or 3 (rarely, e.g. sometimes in Albuca); diplostemonous (usually), or isomerous with the perianth; alterniperianthial (usually), or oppositiperianthial; filantherous (the filaments often broad and flat). Filaments appendiculate (sometimes appendaged by lobes on either side of the anther), or not appendiculate. Anthers dorsifixed (epipeltate); dehiscing via longitudinal slits; introrse; tetrasporangiate; unappendaged. The endothecial thickenings spiral. Microsporogenesis simultaneous. Tapetum glandular. Pollen grains aperturate; 1 aperturate; sulcate; 2-celled.


Gynoecium 3 carpelled. Carpels isomerous with the perianth. The pistil 3 celled. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 3 locular. Gynoecium stylate, or non-stylate to stylate. Styles attenuate from the ovary, or from a depression at the top of the ovary; apical. Stylar canal present. Stigmas 1, or 3; wet type, or dry type; papillate. Placentation axile. Ovules 2–50 per locule; arillate (e.g., Lachenalia), or non-arillate; anatropous; bitegmic; crassinucellate, or pseudocrassinucellate. Embryo-sac development Polygonum-type (usually), or Allium-type, or Scilla-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed, or not formed (then the three nuclei degenerating early); when formed, 3; not proliferating; ephemeral, or persistent. Synergids pear-shaped (with filiform apparatus). Endosperm formation helobial (usually), or nuclear.

Fruit non-fleshy; dehiscent; a capsule. Capsules loculicidal. Seeds endospermic. Endosperm oily. Seeds wingless. Cotyledons 1. Embryo achlorophyllous (5/6); straight (usually), or curved. Testa perhaps always encrusted with phytomelan; black (or sometimes to chestnut in Eucomis).

Seedling. Hypocotyl internode absent. Seedling collar not conspicuous. Cotyledon hyperphyll elongated, or compact; assimilatory, or non-assimilatory; when elongated, more or less circular in t.s. Coleoptile absent. First leaf centric. Primary root ephemeral.

Physiology, biochemistry. Cyanogenic (Albuca). Alkaloids absent. Proanthocyanidins absent. Flavonols present, or absent; when present, kaempferol, or kaempferol and quercetin. Ellagic acid absent. Saponins/sapogenins present (often abundantly). Inulin recorded (Gibbs 1974). C3 physiology recorded directly in Ornithogalum, Scilla. Anatomy non-C4 type (Ornithogalum, Scilla).


Geography, cytology. Holarctic, Paleotropical, Cape, and Antarctic. Patagonian. Widely distributed, richest in southern Africa and from the Mediterranean to southwest Asia.

Taxonomy. Subclass Monocotyledonae. Dahlgren et al. Superorder Liliiflorae; Asparagales. APG 3 core angiosperms; Superorder Lilianae; non-commelinid Monocot; Order Asparagales (as a synonym of Asparagaceae).

Species 500–700. Genera about 30; Albuca, Alrawia, Amphisiphon, Androsiphon, Barnardia, Battandiera, Bellevalia, Brimeura, Bowiea, Camassia, Chionodoxa, Chlorogalum, Daubenya, Dipcadi, Drimia, Drimiopsis, Eucomis, Fortunatia, Galtonia, Hyacynthella, Hyacynthoides (= Endymion), Hyacynthus, Lachenalia, Ledebouria, Leopoldia, Litanthus, Massonia, Muscari, Muscarimia, Neobakeria, Neopatersonia, Ornithogalum, Periboea, Paradisea (or Asphodelaceae, or Anthericaceae?), Polyxena, Pseudogaltonia, Puschkinia, Resnova, Rhadamanthus, Rhodocodon, Schizobasis, Schizocarphus, Schoenolirion, Scilla, Tenicroa, Thuranthos, Urginea, Veltheimia, Whiteheadia.


Illustrations.
• Technical details: Agraphis (= Endymion), Hyacinthus, Muscari).
• Daubenya aurea: Bot. Reg. 1813, 1836.
• Daubenya fulva: Bot. Reg. 1839, 53.
• Drimia villosa: Bot. Reg. 1346, 1830.
• Endymion non-scriptus: ‘Bluebells’ (photos).
• Endymion non-scriptus (B. Ent.).
• Hyacinthus hyacinthoides: as H. spicatus, Bot. Reg. 1869 (1836).
• Lachenalia pallida: Bot. Reg. 1350, 1830.
• Lachenalia, Muscari (Chittenden).
• Massonia candida: Bot. reg. 694, 1823.
• Ornithogalum montanum: Bot. Reg. XXIV, 28 (1838).
• Ornithogalum umbellatum.
• Oenothera anomala: Bot. Mag. 1797.
• Scilla cupani: as S. cupaniana, Bot. Reg 1878 (1836).
• Scilla autumnalis: Eng. Bot. 1526 (1869).
• Scilla verna: Eng. Bot. 1527 (1869).
• Veltheimia, Ornithogalum (Chittenden).

Quotations

The azur’d harebell, like thy veins
(‘Cymbeline’, iv., 2 — re. Endymion non-scriptus, the vulgar names ‘harebell’ and ‘bluebell’ having been transposed in quite recent times. Thus, Shakespeare’s harebells flower in early summer, alongside primroses)

In the lone copse or shadowy dell,
Wild cluster’d knots of harebells blow
(Matthew Robinson’s undated (19th Century) ‘New Family Herbal’)

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