| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Nelumbonaceae Dum.

Habit and leaf form. Aquatic herbs; laticiferous (with articulated laticifers). Without conspicuous aggregations of leaves; rhizomatous. Hydrophytic (with emergent leaves and flowers); rooted. Leaves emergent, or floating. Conspicuously heterophyllous (in the sense that the leaf primordia arise in threes: one giving rise to a scale-leaf on the underside of the rhizome, which at first envelopes the terminal bud, while the second develops into an upper scale leaf which enwraps the petiole base of the large laminate leaf). Leaves medium-sized, or large; with the peculiar arrangement described; foliage leaves long petiolate; simple; peltate. Lamina entire; palmately veined. Leaves stipulate. Stipules intrapetiolar; concrescent; ochreate. Leaf development not ‘graminaceous’.

Leaf anatomy. Stomata present; anomocytic. Lamina with secretory cavities. Secretory cavities containing latex. The mesophyll without sclerenchymatous idioblasts. Foliar vessels absent.

Axial (stem, wood) anatomy. Secretory cavities present; with latex. Primary vascular tissues consisting of scattered bundles. Secondary thickening absent. The axial xylem without vessels.

The axial xylem presumably with tracheids.

Root anatomy. Root xylem with vessels.

Reproductive type, pollination. Unisexual flowers absent. Plants hermaphrodite. Pollination entomophilous; via beetles.

Inflorescence, floral, fruit and seed morphology. Flowers solitary; bracteate (via an upper scale leaf); ebracteolate; large; regular; acyclic. The perianth acyclic and the androecium acyclic, or the perianth acyclic, the androecium acyclic, and the gynoecium acyclic (the gynoecium in ‘more or less distinct’ whorls). Free hypanthium absent.

Perianth with distinct calyx and corolla (the outer several members perhaps interpretable as K), or sequentially intergrading from sepals to petals; about 22–30; free. Calyx (if so interpreted) 2–8; polysepalous; not persistent; imbricate. Corolla (if the inner members so interpreted) 18–28; polypetalous; imbricate; yellow, or red to pink.

Androecium about 200–400. Androecial members maturing centripetally; free of the perianth; free of one another; spiralled. Androecium exclusively of fertile stamens. Stamens about 200–400; filantherous. Anthers adnate; non-versatile; dehiscing via longitudinal slits; introrse and latrorse; tetrasporangiate; appendaged. The anther appendages apical (as a flat or clavate extension of the connective). Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. Tapetum glandular. Pollen shed as single grains. Pollen grains aperturate; 3 aperturate; colpate; 3-celled.

Gynoecium 12–40 carpelled; apocarpous; eu-apocarpous; superior, or inferior (depending on interpretation, the individual carpels being embedded in the spongy receptacle). Carpel apically stigmatic (the stigma annular-dilated); 1(–2) ovuled. Placentation apical to marginal (ventral). Ovules pendulous; apotropous; with dorsal raphe; non-arillate; anatropous; bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Embryo-sac development Polygonum-type. Antipodal cells formed; 3; not proliferating; very ephemeral. Synergids pear-shaped. Endosperm formation cellular. Embryogeny asterad.

Fruit non-fleshy (but the individual carpels sunken in the spongy receptacle); an aggregate. The fruiting carpels not coalescing (ultimately — free within the receptacle, released separately as it disintegrates), or coalescing into a secondary syncarp (i.e. this structure being not unreasonably described as such). The fruiting carpel indehiscent; nucular. Fruit loosely enclosed with its neighbours in the spongy, swollen receptacle, finally released by decay; 1 seeded. Seeds non-endospermic. Perisperm absent. Cotyledons 2 (connate, and sheathing the plumule). Embryo chlorophyllous (1/2).

Physiology, phytochemistry. Not cyanogenic. Alkaloids present. Iridoids not detected. Saponins/sapogenins absent. Proanthocyanidins present. Ellagic acid absent. Sieve-tube plastids S-type.

Geography, cytology. Sub-tropical to tropical. Eastern U.S.A. to Colombia, and warm Asia to Northeast Australia. X = 16.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Magnoliiflorae; Nelumbonales. Cronquist’s Subclass Magnoliidae; Nymphaeales. APG III core angiosperms; peripheral eudicot; Superorder Proteanae. APG IV Order Proteales.

Species 2 (accounted for here). Genera 1 (accounted for here); the only known surviving genus being Nelumbo, with others (e.g., Nelumbites, Exnelumbites, Paleonelumbo, Nelumbago, and Notocyamus) known only from fossils.

Economic uses, etc. N. pentapetala rhizomes provided starchy food for American Indians.

Illustrations. • Le Maout and Decaisne: Nelumbo lutea (Lotus). • Nelumbo lutea (as Nelumbium): Bot Mag. 66 (1839). • Nelumbo nucifera (as Nelumbium speciosum): Lindley. • Nelumbo nucifera var. caspica: as Nelumbium caspicum, Bot. Reg. 1844, 14. • Nelumbo nucifera (as Nelumbium speciosum): R. Wight (1840).

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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