Flora Mesoamericana, Vol. 2 (1), Nelumbonaceae, page 1 of 3

First published on the Flora Mesoamericana Website, 30 Mar. 2015.

57A. NELUMBONACEAE By G. Davidse.

Perennial, herbaceous, rhizomatous, aquatic herbs with white latex; air chambers conspicuous in vegetative portions of plant. Rhizomes long and branching, rooting at the nodes, producing elongated tubers at the tips late in the growing season. Leaves arising directly from the rhizomes, spirally and alternately arranged, long-petiolate, peltate, the earliest ones floating, the later ones long-emergent from the water, the vernation involute; petioles circular in cross-section; blades nearly circular, with a central disc that connects to the apex of the petiole, the margins entire, the main veins radiating from the central disc, dichotomously branched in approximately the upper 1/3, the ultimate veinlets loop- connected just below the blade margin. Inflorescences reduced to solitary, axillary flowers. Flowers large, bisexual, actinonomorphic, hypogynous, protogynous, diurnal, long-peduculate, well emergent from the water and normally exceeding the leaves; tepals numerous, distinct, the outermost reduced, sepaloid, the inner tepals becoming larger and more petaloid; stamens numerous, the filaments slender; the anthers long and narrow, dehiscing by longitudinal slits, with prominent, incurved, connective appendages, these thicker than the fertile portion and lighter in color; pistils numerous, 1-carpellate embedded separately in the cavities of an enlarged, turbinate receptacle with only the styles and stigmas protruding, the ovary 1-locular, the placentation apical, the ovule 1, the style short, the stigma nearly sessile, capitate with a central depression; fruiting receptacle much enlarged, broadly obovate in longitudinal section, the top flattened, the walls smooth, prominently fluted. Fruits nutlike, indehiscent, hard, smooth, embedded but loose in the cavities of the enlarged receptacle; seed 1, without endosperm, the embryo completely filling the seed. 1 gen., 2 spp. Native to eastern North America and eastern Asia to Australia; introduced into other tropical and subtropical areas. The genus Nelumbo has been traditionally treated as part of the Nymphaeaceae, but molecular data clearly shows that is not closely related to that family and surprisingly Flora Mesoamericana, Vol. 2 (1), Nelumbonaceae, page 2 of 3 falls into the Proteales and should be recognized in its own family (summarized in Stevens, 2001 onwards). The two species have been used in many ways in various human societies, including for food and medicinal purposes, but perhaps most famously as a sacred symbol in Buddhism and Hinduism. Incredibly, seeds of the Old World Nelumbo nucifera Gaertn. have been shown to maintain viability for 1300 years (Shen-Miller et al., 1995, 2002; Borsch & Barthlott, 1994; Yatskievych, 2013). They are today widely cultivated as aquatic ornamentals. Bibliography: Wiersema, J. H. Fl. N. Amer. 3: 64-65 (1993).

1. Nelumbo Adans. Nelumbium Juss. Por G. Davidse. See family description.

1. Nelumbo lutea Willd., Sp. Pl. 2: 1259 (1799). Neotype (designated by Wiersema & Reveal, 1991): United States, Bartram s.n. (BM!). Illustr.: Yatskievych, 2013: 475, t. 458g-h. Nelumbium luteum Willd., Nelumbo nucifera Gaertn. subsp. lutea (Willd.) Borsch & Barthlott, N. nucifera var. lutea (Willd.) Kuntze, N. pentapetala (Walter) Fernald, Nymphaea pentapetala Walter. Tubers 5-15 cm long. Leaves 75-300 cm; petioles smooth or rarely with a few prickles; blades 20-60(-80) cm in diameter. Peduncles smooth or rarely with a few prickles. Flowers to 24 cm in diameter, pale yellow to nearly white; tepals 14-30, 7-12 cm, the outermost 1-5, shorter, normally long persistent and reflexed during fruit development, greenish on the back; stamens 100-200, 2.5-3.0 cm, the anthers 1-2 cm, the sterile appendages 5-7 mm. Receptacle with 8-32 embedded pistils, enlarging to 10 cm in diameter at fruiting maturity, ultimately bending downwards to release the seeds through enlarged openings. Fruits 9-16 × 8-13 mm, nearly globose to slightly elongated at maturity. 2n = 16. Edges of ponds and lakes in stagnant water. 0-800 m. T (Novelo y Flora Mesoamericana, Vol. 2 (1), Nelumbonaceae, page 3 of 3

Ramos 2311, MO); C (Ocaña y Vera 70, MO); H (Chorley y Atkinson 80, MO ). Native to eastern North America; probably introduced and naturalized elsewhere.) This species is closely related to the Old World sacred lotus, Nelumbo nucifera Gaertn, and was treated as a subspecies of it by Borsch & Barthlott (1994). Nelumbo nucifera has pink flowers (vs. pale yellow) and the petioles have numerous prickles (vs. smooth or nearly so).

Bibliography

Borsch, T. & W.A. Barthlott. 1994. Classification and distribution of the genus Nelumbo Adans. (Nelumbonaceae). Beitr. Biol. Pflanzen 68: 421–450. Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 13, September 2013 [and more or less continuously updated since]. http://www.mobot.org/MOBOT/research/APweb/. Wiersema, J. H. 1993. Nelumboaceae. Fl. N. Amer. 3: 64-65. Yatskievych, G. 2013. Nelumbonaceae. In G. Yatskievych, Steyermark's Fl. Missouri 3: 479–481. The Missouri Botanical Garden Press, St. Louis.