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The families of flowering plants

L. Watson and M.J. Dallwitz

Moringaceae Dum.

Including Hyperanthereae (Hyperantheraceae) Link

Habit and leaf form. Trees (of Acacia-like habit, commonly stout-trunked with gummy bark, and with a large mucilage canal in the centre of the pith); resinous. Leaves deciduous (with deciduous leaflets); alternate; petiolate; non-sheathing; compound; bipinnate to multiply compound (2–3 times pinnate, with opposite leaflets). Lamina cross-venulate. Leaves stipulate, or exstipulate. Stipules when present represented by glands (and similar structures present at the bases of the pinnae).

Leaf anatomy. The leaf lamina dorsiventral. Mucilaginous epidermis often present, or absent. Stomata present; anomocytic. Hairs present; glandular; unicellular (with blunt tips). Unicellular hairs branched. Complex hairs absent. The mesophyll containing crystals (as well as abundant myrosin cells). The crystals druses.

Axial (stem, wood) anatomy. Secretory cavities present (in the pith, a large central one sometimes accompanied by a second, smaller one); with mucilage. Primary vascular tissues in a cylinder, without separate bundles; collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring. Primary medullary rays narrow.

The wood ring porous. The vessels medium sized, commonly in radial multiples. The vessel end-walls simple. The vessels with vestured pits; without spiral thickening. The axial xylem without fibre tracheids; with libriform fibres; without septate fibres. The fibres without spiral thickening. The parenchyma paratracheal (vasicentric or slightly aliform). The secondary phloem not stratified. ‘Included’ phloem absent. The wood storied.

Reproductive type, pollination. Plants hermaphrodite.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in panicles. The ultimate inflorescence units cymose. Inflorescences axillary; many-flowered axillary panicles. Flowers somewhat irregular, or very irregular; zygomorphic. The floral irregularity involving the perianth and involving the androecium. Flowers 5 merous; cyclic; pentacyclic. Floral receptacle developing a gynophore. Free hypanthium present (short). Hypogynous disk present (lining the hypanthium); intrastaminal; annular (with a short free margin).

Perianth with distinct calyx and corolla (but the calyx petaloid); 10; 2 whorled; isomerous. Calyx 5 (petaloid); 1 whorled; polysepalous (the sepals spreading or reflexed); unequal but not bilabiate (the sepals unequal); imbricate; with the median member posterior. Corolla 5; 1 whorled; polypetalous; imbricate; unequal but not bilabiate (with the outer, anterior member the largest, the two inner (upper, posterior) members reflexed and the smallest, the lateral pair ascending); white, or red.

Androecium (7–)10. Androecial members free of the perianth (inserted at the mouth of the hypanthium, around the disk); markedly unequal (declinate); free of one another; 2 whorled. Androecium including staminodes. Staminodes (3–)5; external to the fertile stamens (the outer, antesepalous whorl); non-petaloid (setiform or resembling antherless filaments). Stamens 5; isomerous with the perianth; oppositisepalous; both alternating with and opposite the corolla members; filantherous (the filaments sometimes slightly petaloid). Anthers declinate; dorsifixed; dehiscing via longitudinal slits; ultimately unilocular; bisporangiate (mostly), or tetrasporangiate (rarely). Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Tapetum glandular. Pollen shed as single grains. Pollen grains aperturate; (2–)3(–4) aperturate; colporate; 2-celled.

Gynoecium 3 carpelled. Carpels reduced in number relative to the perianth. The pistil 1 celled. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 1 locular; stipitate (curved). Gynoecium stylate. Styles 1 (recurved); apical. Stylar canal present. Stigmas 1; truncate. Placentation parietal. Ovules in the single cavity 40–100 (‘many’); pendulous; biseriate; non-arillate; anatropous (the radicle superior); bitegmic; crassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing only after one has been fertilized, or fusing simultaneously with the male gamete (?). Antipodal cells formed; 3; not proliferating; very ephemeral. Synergids hooked (with faint filiform apparatus). Endosperm formation nuclear. Embryogeny asterad.

Fruit non-fleshy; dehiscent; a capsule (large, woody, 3–12 angled, unilocular, podlike, torulose, but without a replum). Capsules three valvular (the seeds attached medially along the valves). Fruit elastically dehiscent (explosively). Seeds non-endospermic; large; three winged (usually), or wingless. Embryo well differentiated. Cotyledons 2. Embryo chlorophyllous (1/1); straight. Micropyle zigzag.

Physiology, phytochemistry. Sugars transported as sucrose. Mustard-oils present (with myrosin cells abundant in leaves, stems and roots). Not cyanogenic. Alkaloids present, or absent. Iridoids not detected. Saponins/sapogenins absent. Proanthocyanidins absent. Flavonols present; kaempferol and quercetin. Ellagic acid absent. Aluminium accumulation not found.

Geography, cytology. Holarctic and Paleotropical. Temperate to tropical. Mediterranean and North Africa to Arabia and India, also Southwest africa and Madagascar. X = 14.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Violiflorae; Capparales. Cronquist’s Subclass Dilleniidae; Capparales. APG III core angiosperms; core eudicot; Superorder Rosanae; malvid. APG IV Order Brassicales.

Species 12, or 13. Genera 1; only genus, Moringa.

Economic uses, etc. Moringa oleifera (horseradish tree) is cultivated throughout the tropics as a valuable, multi-purpose source of food for people and livestock. It has edible fruits, the leaves are rich in proteins with essential amino acids, the seeds supply ‘oil of ben’ used in perfumery and light lubricants, and the roots are a source of edible condiment. The similar potential of other species has as yet been less extensively exploited.

Illustrations. • Le Maout and Decaisne: Moringa. • Moringa pterygosperma (cf. oleifera): R. Wight (1840). • Moringa concanensis: Hook. Ic. Pl. 26 (1899).

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: 5th March 2018.’.