Nymphaeaceae Salisb. Including Nupharaceae Nak.Excluding Barclayaceae, Cabombaceae, Euryalaceae Habit and leaf form. Aquatic herbs; laticiferous.

Perennial; rhizomatous. Hydrophytic; rooted. Leaves floating. Leaves medium-sized to large; alternate; spiral; petiolate; simple; usually more or less peltate. Lamina entire; usually more or less orbicular; basically palmately veined; cross-venulate; usually cordate. Leaves stipulate (the stipules median-axillary), or exstipulate; without a persistent basal meristem. General anatomy. Plants with laticifers (articulated). The laticifers in leaves, in stems, in roots, in flowers, and in the fruits. Accumulated starch other than exclusively pteridophyte type. Leaf anatomy. Stomata present; anomocytic. The mesophyll with sclerencymatous idioblasts. Minor leaf veins without phloem transfer cells (Nymphaea). Vessels absent. Stem anatomy. Primary vascular tissue comprising a ring of bundles, or in scattered bundles. Secondary thickening absent. Xylem presumably with tracheids; without vessels. Sieve-tube plastids S-type. Pith with diaphragms, or without diaphragms. Root anatomy. Root xylem without vessels. Reproductive type, pollination. Plants hermaphrodite. Pollination entomophilous. Inflorescence, floral, fruit and seed morphology. Flowers solitary; large; often fragrant; regular; partially acyclic. The perianth acyclic and the androecium acyclic. Free hypanthium absent. Perianth with distinct calyx and corolla (e.g. Nymphaea, according to the usual interpretation), or petaline (Nuphar); 5, or 2050 (many); free. Calyx as commonly interpreted 4, or 5; polysepalous; imbricate. Corolla 5 (Nuphar), or 1550 (many, showy); polypetalous; imbricate. Androecium 4080. Androecial members maturing centripetally; free of the perianth; free of one another; spiralled. Androecium including staminodes (with transition from petals to stamens). Staminodes 1120 (in the form of nectariferous scales, in Nuphar); external to the fertile stamens; in Nuphar, petaloid. Stamens 4080; petaloid, or laminar, or filantherous (in sequence). Anthers adnate; non-versatile; dehiscing via longitudinal slits; introrse; locule number?; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with more than one middle layer. Tapetum glandular. Pollen shed as single grains. Pollen grains aperturate; 1 aperturate; sulcate, or zoniaperturate; 2-celled. Gynoecium 535 carpelled. The pistil 535 celled. Gynoecium syncarpous; eu-syncarpous; superior to partly inferior. Ovary 535 locular. Placentation more or less parietal (or ovules more or less scattered). Ovules 10100 per locule (i.e. many); arillate, or non-arillate; orthotropous, or 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 fleshy; dehiscent, or indehiscent, or a schizocarp (irregularly so, Nuphar). Mericarps in Nuphar, 535 (?). Fruit when non-schizocarpic, a berry; enclosed in the fleshy receptacle. Seeds endospermic. Perisperm present. Cotyledons 1, or 2. Embryo chlorophyllous (2/2); straight. Seedling. Germination cryptocotylar. Physiology, biochemistry. Not cyanogenic. Alkaloids present (usually), or absent. Iridoids not detected. Proanthocyanidins present, or absent; cyanidin, or delphinidin. Flavonols present, or absent (Nuphar); in Nymphaea, kaempferol and quercetin, or myricetin. Ellagic acid present (one Nymphaea sample), or absent (Nymphaea capensis, Nuphar lutea). Saponins/sapogenins absent. Aluminium accumulation not found. C 3. C3 physiology recorded directly in Nuphar. Anatomy non-C4 type (Brasenia, Nuphar, Nymphaea). Geography, cytology. Temperate to tropical. Cosmopolitan, except in frigid zones. Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgrens Superorder Nymphaeiflorae; Nymphaeales. Cronquists Subclass Magnoliidae; Nymphaeales. APG 3 peripheral angiosperms; Superorder Nymphaeanae; Order Nymphaeales. Species 75. Genera 3; Nymphaea, Nuphar, Ondinea.

K 4-6 [-14] C 8- [0] A G (3-), superior or inferior.

Quotations The Water Lilies, white and yellow flowers, How beautiful they are upon the lake! (John Clare 1841, The Water Lilies Nymphaea alba) The Nymphaeaceae are aquatic, rhizomatous herbs comprising eight genera and 70 species. The family is further characterized by scattered vascular bundles in the stems, and frequent presence of latex. Leaves mostly alternate, simple, entire to dissected, submerged, floating, or emergent; stipules present or absent. Flowers solitary, bisexual, actinomorphic, long-pedicellate, floating or emergent; perianth of 4-6-many, commonly poorly differentiated, distinct or connate, often petaloid segments, these often blending with staminodes. Androecium of 3-many, laminar to filamentous stamens and commonly numerous petaloid or reduced staminodes, including transitional forms. Gynoecium apocarpous or more often syncarpous, of 3-numerous carpels, syncarpous forms with as many locules as carpels and with an equal number of elongated, radiating stigmas; ovary superior to inferior; placentation parietal; ovules 1-numerous (commonly scattered on the partitions). Fruit an aggregate of nuts, a berry, or an irregularly dehiscent fleshy capsule. Seeds often arillate, more or less lacking endosperm but with abundant perisperm.

Illiciaceae Van Tiegh. ~ Schisandraceae Habit and leaf form. Glabrous, small trees, or shrubs; bearing essential oils. Leaves evergreen; alternate; spiral (sometimes crowded towards the tips of the twigs); leathery; petiolate; non-sheathing; gland-dotted, or not glanddotted; aromatic, or without marked odour; simple. Lamina entire; pinnately veined. Leaves exstipulate. Lamina margins entire. Leaf anatomy. Stomata present; mainly confined to one surface (abaxial); anomocytic (occasionally), or paracytic (usually). Lamina dorsiventral. The mesophyll with spherical etherial oil cells. Minor leaf veins without phloem transfer cells. Stem anatomy. Nodes unilacunar (with one trace). Internal phloem absent. Secondary thickening developing from a conventional cambial ring. Included phloem absent. Xylem with tracheids; with vessels. Vessel end-walls scalariform and reticulately perforated. Primary medullary rays narrow. Wood parenchyma paratracheal. Sievetube plastids S-type. Reproductive type, pollination. Plants hermaphrodite. Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in inflorescences (these two- or three-flowered). Inflorescences axillary (or supra-axillary). Flowers small; regular; partially acyclic. The perianth acyclic and the androecium acyclic (the perianth and stamens commonly arranged in several series, but spiral within each). Free hypanthium absent. Hypogynous disk absent. Perianth sequentially intergrading from sepals to petals (the members of the outermost series small and bractlike or sepaloid, the inner members commonly larger and more petaloid), or sepaline (the inner members reduced and sometimes transitional with the stamens); 733; free; commonly in several series, spiral within each. Androecium (4)2050. Androecial members maturing centripetally (?); free of the perianth; free of one another; spirally arranged, in (1)several series. Androecium exclusively of fertile stamens, or including staminodes. Staminodes external to the fertile stamens ((when present) in the form of intermediates with the inner perianth members). Stamens (4)1550; filantherous (the filaments short and thick). Anthers adnate (the thecae lateral); non-versatile; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral. Anther wall initially with more than one middle layer. Tapetum glandular. Pollen shed as single grains. Pollen grains aperturate; 3 aperturate; colpate (colpoidate); 2-celled. Gynoecium (5)715(21) carpelled; apocarpous; eu-apocarpous (its members attached obliquely to the receptacle in a single cycle, this ontogenetically resolvable into a tight helix); superior. Carpel incompletely closed (being unsealed at the narrowed, stylar tip); stylate; with a decurrent stigma; 1 ovuled. Placentation ventral and nearly basal. Stigmas dry type; non-papillate. Ovules ascending; anatropous; bitegmic; crassinucellate. Embryo-sac development Polygonum-type. Antipodal cells formed; 3; not proliferating; ephemeral. Endosperm formation cellular, or nuclear. Embryogeny asterad. Fruit non-fleshy; an aggregate. The fruiting carpel dehiscent; a follicle (the cycle of follicles often spreading radially in a stellate pattern). Fruit 1 seeded. Seeds copiously endospermic. Endosperm oily. Embryo well differentiated (very small). Cotyledons 2. Embryo achlorophyllous (1/1). Seedling. Germination phanerocotylar. Physiology, biochemistry. Not cyanogenic. Iridoids not detected. Proanthocyanidins present; cyanidin and delphinidin. Flavonols present; kaempferol and quercetin. Ellagic acid absent. Geography, cytology. Holarctic, Paleotropical, and Neotropical. Sub-tropical to tropical. Tropical Southeast Asia, North America, West Indies. X = 13, 14.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgrens Superorder Magnoliiflorae; Illiciales. Cronquists Subclass Magnoliidae; Illiciales. APG 3 peripheral angiosperms; Superorder Austrobaileyanae; Order Austrobaileyales (as a synonym of Schisandraceae?). Species 42. Genera 1; only genus, Illicium. Economic uses, etc. Some cultivated ornamentals, and commercial volatile oil from I. verum.

Chloranthaceae R. Br. Habit and leaf form. Trees, shrubs, and herbs; bearing essential oils. Annual, or perennial. Leaves opposite; petiolate; connate; aromatic; simple; epulvinate; stipulate. Stipules interpetiolar. Lamina margins serrate. Leaves without a persistent basal meristem. General anatomy. Accumulated starch other than exclusively pteridophyte type. Leaf anatomy. Adaxial hypodermis commonly present. Lamina dorsiventral; with secretory cavities, or without secretory cavities. Secretory cavities when present, containing mucilage. The mesophyll with spherical etherial oil cells; without calcium oxalate crystals. Minor leaf veins without phloem transfer cells (Chloranthus). Stem anatomy. Stems with solid internodes. Young stems usually cylindrical. Secretory cavities present, or absent; when present, with mucilage. Nodes unilacunar (clearly, in some genera), or tri-lacunar (or interpretable as a modification of such, in some genera). Secondary thickening developing from a conventional cambial ring. Xylem with fibre tracheids, or without fibre tracheids; with libriform fibres, or without libriform fibres; with vessels, or without vessels. Vessel end-walls oblique; scalariform. Wood parenchyma paratracheal. Sieve-tube plastids S-type. Reproductive type, pollination. Plants hermaphrodite, or dioecious. Inflorescence, floral, fruit and seed morphology. Flowers aggregated in inflorescences; in spikes, or in heads, or in panicles. The ultimate inflorescence unit cymose. Inflorescences terminal, or axillary; spikes or cymes. Flowers two bracteolate; small; reduced. Perianth sepaline (female and hermaphrodite flowers), or absent (male flowers); when present, 3; when present, joined; when present, 1 whorled. Androecium 15. Androecial members free of the perianth; united with the gynoecium; usually more than one, coherent; 1 adelphous (more or less connate). Androecium exclusively of fertile stamens, or including staminodes (or at least the laterals sometimes with only half-anthers). Stamens 13(5); laminar, or filantherous. Anthers dehiscing via longitudinal slits, or dehiscing by longitudinal valves; unilocular to bilocular; tetrasporangiate, or bisporangiate and tetrasporangiate (in Chloranthus). Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral. Anther wall initially with more than one middle layer. Pollen grains aperturate, or nonaperturate; 16 aperturate; sulcate, or colpate, or porate; 2-celled. Gynoecium when present, 1 carpelled. The pistil 1 celled. Gynoecium monomerous; of one carpel; superior, or partly inferior. Carpel 1 ovuled. Placentation apical. Stigmas dry type; non-papillate; Group II type. Ovules pendulous; anatropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle, or not contributing to the micropyle. Embryo-sac development Polygonum-type. Antipodal cells formed; initially 3; proliferating (sometimes, up to 40 cells), or not proliferating. Synergids pear-shaped. Endosperm formation cellular. Embryogeny onagrad, or chenopodiad. Fruit fleshy. The fruiting carpel indehiscent; drupaceous. Seeds endospermic. Endosperm oily (and starchy). Perisperm present. Seeds with starch. Embryo rudimentary at the time of seed release to well differentiated (very small). Cotyledons when differentiated, 2. Physiology, biochemistry. Proanthocyanidins absent. Flavonols absent. Ellagic acid absent (Chloranthus). Aluminium accumulation not found. Geography, cytology. Sub-tropical to tropical. South and Central America, Southeast Asia and Malaysia. X = 8, 14, 15. Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgrens Superorder Magnoliiflorae; Magnoliales. Cronquists Subclass Magnoliidae; Piperales. APG 3 core angiosperms; unplaced at Superordinal level; Order Chloranthales. Species 65. Genera 4; Ascarina, (Ascarinopsis), Chloranthus, Hedyosmum, Sarcandra. General remarks. See Leroy 1983.