| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Melanthiaceae Batsch

~ Former Liliaceae.

Including Abaminaceae J.G. Agardh, Chionographidaceae, Cymbantheae (Cymbanthaceae) Salisb.(?), Heloniadaceae J.G. Agardh, Miyoshiaceae Makino, Protoliriaceae Makino, Veratraceae von Vest; excluding Nartheciaceae, Petrosaviaceae

Habit and leaf form. Herbs. Leaves well developed. Plants non-succulent. Annual (rarely), or perennial; with a basal aggregation of leaves, or without conspicuous aggregations of leaves; rhizomatous (mainly), or tuberous, or cormous (rarely). Leaves alternate; spiral (usually), or distichous; flat; ‘herbaceous’; sessile (usually), or petiolate (occasionally, almost); sheathing. Leaf sheaths not tubular. Leaves with blades borne edgewise to the stem, or with blades ‘normally orientated’; simple. Lamina entire; linear to lanceolate (usually), or ovate (rarely); parallel-veined. Leaves eligulate.

Leaf anatomy. The leaf lamina dorsiventral. The mesophyll commonly containing crystals. The crystals raphides, or solitary-prismatic. Foliar vessels absent.

Axial (stem, wood) anatomy. Secondary thickening absent. The axial xylem without vessels.

Root anatomy. Root xylem with vessels, or without vessels (in achlorophyllous forms); vessel end-walls scalariform.

Reproductive type, pollination. Unisexual flowers present (rarely), or absent. Plants hermaphrodite (nearly always), or dioecious, or polygamomonoecious. Floral nectaries present. Nectar secretion from the gynoecium (Zigadenus etc. with septal nectaries).

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in racemes, or in spikes, or in panicles. Inflorescences scapiflorous, or not scapiflorous; simple or compound racemes or spikes, rarely panicles. Flowers bracteate, or ebracteate; regular (nearly always), or very irregular (only in Chionographis); in Chionographis, zygomorphic; 3 merous; cyclic; usually pentacyclic. Perigone tube present (in hemi-epigynous forms), or absent. Hypogynous disk absent.

Perianth of ‘tepals’; 6; free to joined; 2 whorled (3+3); isomerous; sepaloid, or petaloid; without spots, or spotted (occasionally); similar in the two whorls; white, or brown, or purple (usually inconspicuous, usually lacking patterns and spurs).

Androecium 6, or 9 (Pleea), or 12 (Pleea). Androecial members more or less free of the perianth; free of one another; 2 whorled (usually 3+3, Pleea with 6+3 or 6+6). Androecium exclusively of fertile stamens. Stamens 6 (usually); diplostemonous (usually); normally alterniperianthial. Anthers dorsifixed (hypopeltate), or basifixed; dehiscing via longitudinal slits; extrorse (usually), or introrse (occasionally). Endothecium developing fibrous thickenings. The endothecial thickenings spiral. Microsporogenesis successive (usually), or simultaneous (Tofieldia). Anther wall initially with more than one middle layer; of the ‘monocot’ type. Tapetum glandular. Pollen grains aperturate; 1 aperturate (usually), or 2 aperturate, or 4 aperturate; sulcate (usually), or sulculate, or foraminate; 2-celled.

Gynoecium 3 carpelled. Carpels isomerous with the perianth. The pistil 3 celled (with Petrosaviaceae excluded). Gynoecium always more or less syncarpous; semicarpous (the carpels often free above), or synovarious (usually, with separate styles), or synstylovarious (Aletris, with a tribrachiate style); superior, or partly inferior. Carpel 2–100 ovuled (to ‘many’). Placentation marginal. Ovary when syncarpous, 3 locular; sessile. Gynoecium stylate. Styles 1, or 3; when three, free (Helonias, Heloniopsis), or partially joined; apical. Stigmas 1, or 3; of Veratrum dry type. Placentation axile. Ovules 2–100 per locule (to ‘many’); funicled; non-arillate; at least, usually anatropous; bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Endothelium differentiated. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; proliferating (Veratrum). Endosperm formation helobial (by contrast with Liliales).

Fruit non-fleshy; an aggregate, or not an aggregate. The fruiting carpel when semicarpous, dehiscent; a follicle (opening along the inner suture). Fruit when syncarpous, dehiscent; a capsule. Capsules septicidal, or loculicidal (or opening ventricidally from apex to base, when the carpels are distally free, e.g. Tofieldia, Veratrieae). Seeds endospermic. Endosperm oily. Seeds usually winged (or with terminal appendages), or wingless. Embryo well differentiated (but often small). Embryo ovoid or globose. Testa without phytomelan (by conrast with most capsular Asparagales, and also lacking phlobaphene).

Seedling. Hypocotyl internode present, or absent. Seedling collar not conspicuous. Cotyledon hyperphyll elongated; assimilatory; more or less circular in t.s., or dorsiventrally flattened (variously unifacial or bifacial). Coleoptile absent. Seedling cataphylls absent. First leaf ensiform (e.g. Tolfieldia), or dorsiventral (e.g. Zigadenus). Primary root ephemeral.

Physiology, phytochemistry. Inulin recorded (Gibbs 1974). Not cyanogenic. Alkaloids present (Veratreae), or absent. Saponins/sapogenins often present. Proanthocyanidins present, or absent; when present, cyanidin. Flavonols present, or absent; when present, kaempferol and quercetin. Ellagic acid absent.

Geography, cytology. Holarctic (mainly), Paleotropical, Neotropical. Frigid zone, temperate (mainly), sub-tropical, tropical. Widespread N. temperate, E. Asia, Malaya, Borneo, the Americas, Arctic — not Africa, not Australia. Chromosomes with diffuse centromeres (Chionographis), or with normal centromeres. X = 8, 11, 12, 13, 15, 17.

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

Species about 140. Genera about 25; Amianthium, Anticlea, Chamaelirium, Chionographis, Helonias, Heloniopsis, Isidrogalvia, Melanthium, Protolirion, Schoenocaulon, Stenanthium, Toxicoscordion, Veratrum, Xerophyllum, Ypsilandra, Zigadenus.

General remarks. An excellent treatment of Melanthiaceae sensu lato by Zomlefer (1997) has yet to be accounted for here.

Illustrations. • Amianthium muscitoxicum: Addisonia 20 (1937–8). • Anticlea elegans var. glaucus, as Zigadenus glaucus: Bot. Reg. XXIV, 67 (1838). Anticlea elegans, as Zigadenus. 1, a single perianth member, with a double gland adaxially at its base. 2, the gynoecium. 3, longitudinally sectioned gynoecium, showing placentation. • Le Maout and Decaisne: Veratrum, Tofieldia, Amianthium (= Zigadenus). • Chionographis japonica: Bot. Mag. 106 (1880). • Helonias bullata: Addisonia 13 (1928). • Heloniopsis orientalis var. breviscapa (as H. japonica): Bot. Mag. 114 (1888). • Schoenocaulon officinale: Köhler’s Medizinal-Pflanzen 2 (1890). • Schoenocaulon officinale: as Sabadilla officinalis, Bot. Reg. 1839, 33. • Veratrum album: Köhler’s Med. Pflanzen 2 (1890).

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: 8th June 2026. delta-intkey.com’.

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