Family Moringaceae

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. Mucilaginous epidermis often present. Stomata present; anomocytic. Hairs present; unicellular.

The mesophyll containing calcium oxalate crystals. The mesophyll crystals druses.

Stem anatomy. Secretory cavities present (in the pith); with mucilage. Internal phloem absent. Secondary thickening developing from a conventional cambial ring. ‘Included’ phloem absent. Xylem without fibre tracheids; with libriform fibres. Vessel end-walls simple. Vessels with vestured pits. Primary medullary rays narrow. Wood ring porous; storied; parenchyma paratracheal (vasicentric or slightly aliform).

Reproductive type, pollination. Plants hermaphrodite.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; in panicles. The ultimate inflorescence unit 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, biochemistry. Mustard-oils present (in myrosin cells). Not cyanogenic. Alkaloids present, or absent. Iridoids not detected. Proanthocyanidins absent. Flavonols present; kaempferol and quercetin. Ellagic acid absent. Saponins/sapogenins absent. Aluminium accumulation not found. Sugars transported as sucrose.

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 3 core angiosperms; core eudicot; Superorder Rosanae; malvid; Order Brassicales.

Species 12. Genera 1; only genus, Moringa.

Economic uses, etc. Moringa oleifera (horseradish tree) has edible fruits, the seeds supply ‘oil of ben’ used in perfumery and light lubricants, and the roots are a source of edible condiment.

Illustrations.
• Technical details: Moringa.