| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Schisandraceae Bl.

Excluding Illiciaceae

Habit and leaf form. Lianas; bearing essential oils. With terminal aggregations of leaves, or without conspicuous aggregations of leaves. Climbing; stem twiners, or scrambling; Schisandra twining clockwise. Leaves persistent, or deciduous; alternate; spiral; ‘herbaceous’, or leathery; petiolate; non-sheathing; gland-dotted (often), or not gland-dotted; aromatic; simple. Lamina entire; pinnately veined; cross-venulate. Leaves exstipulate. Lamina margins entire, or dentate (more often, or at least denticulate).

Leaf anatomy. Mucilaginous epidermis present. Stomata present; mixed anomocytic and paracytic (and ‘laterocytic’). Hairs present, or absent. The mesophyll with spherical etherial oil cells; usually containing mucilage cells. Minor leaf veins without phloem transfer cells (Schisandra).

Axial (stem, wood) anatomy. Young stems with solid internodes. Pith more or less homogeneous (the central cells thinner walled). Cork cambium present; initially superficial. Nodes unilacunar (with three traces). Primary vascular tissues collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring. Primary medullary rays narrow.

The wood diffuse porous. The vessels large; solitary. The vessel end-walls scalariform, or simple. The vessels with spiral thickening, or without spiral thickening (?). The axial xylem with tracheids; without fibre tracheids; without libriform fibres. The parenchyma apotracheal (terminal only). ‘Included’ phloem absent. The wood not storied. Tyloses present.

Reproductive type, pollination. Unisexual flowers present. Plants monoecious, or dioecious.

Inflorescence, floral, fruit and seed morphology. Flowers solitary, or aggregated in ‘inflorescences’ (these two- to few-flowered). Inflorescences axillary, or cauliflorous (occasionally). Flowers bracteolate, or ebracteolate; small; usually fragrant; regular; acyclic. The perianth acyclic, the androecium acyclic, and the gynoecium acyclic (i.e., spiralled throughout). Floral receptacle not markedly hollowed (conic to cylindrical or obovoid). Free hypanthium absent. Hypogynous disk absent.

Perianth sequentially intergrading from sepals to petals, or of ‘tepals’; (5–)9–15(–24) (the outermost and innermost members sometimes more or less reduced); free; spiralled, in two to several series; sepaloid, or petaloid, or sepaloid and petaloid; green (the outer members, often), or white, or cream, or yellow, or red, or pink.

Androecium of male flowers, 4–80. Androecial members maturing centripetally; free of the perianth; free of one another to coherent (the filaments connate basally to wholly connate in a globular, fleshy mass); spiralled. Androecium exclusively of fertile stamens, or including staminodes (in some Kadsura species). Staminodes when present, 3–22; internal to the fertile stamens (i.e., above the fertile stamens in the male flowers); non-petaloid (subulate, sometimes with vestigial thecae). Stamens 4–80; filantherous (the filaments short). Anthers basifixed, or adnate (with separate thecae); non-versatile; dehiscing via longitudinal slits; extrorse to introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or decussate, or tetrahedral and decussate. Anther wall initially with one middle layer, or initially with more than one middle layer (1 to 3); of the ‘dicot’ type. Tapetum glandular. Pollen shed as single grains. Pollen grains aperturate; 6 aperturate (typically), or 3 aperturate (rarely — usually with three short colpi alternating with three long ones, the latter meeting at one pole); colpate (to distally syncolpate); 2-celled.

Gynoecium 12–300 carpelled; apocarpous; eu-apocarpous (spirally arranged); superior. Carpel incompletely closed (unsealed); non-stylate (with a stigmatic crest, which in Kadsura forms a ‘pseudostyle' or a ‘pseudostigma'); with a longitudinal stigmatic surface (the stigma decurrent); (1–)2–5(–11) ovuled. Placentation marginal. Stigmas wet type; papillate; Group III type. Ovules pendulous; collateral, or superposed; anatropous to campylotropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; very ephemeral. Synergids pear-shaped. Endosperm formation cellular. Embryogeny onagrad, or asterad.

Fruit fleshy; an aggregate (the fruiting carpels on a somewhat elongate axis, or in a dense head). The fruiting carpel indehiscent; baccate. Fruit typically 2 seeded. Seeds copiously endospermic. Endosperm oily (and starchy). Seeds flattened. Seeds with starch. Embryo well differentiated (but minute). Cotyledons 2. Embryo straight.

Physiology, phytochemistry. Not cyanogenic. Arbutin absent. Iridoids not detected. Saponins/sapogenins absent. Proanthocyanidins present; cyanidin and delphinidin. Flavonols present; kaempferol, quercetin, and myricetin. Ellagic acid absent (Kadsura). Sieve-tube plastids S-type.

Geography, cytology. Holarctic and Paleotropical. Temperate to tropical. Eastern and Southeastern Eurasia, Malaysia, Southeastern U.S.A. X = 13, 14.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Magnoliiflorae; Illiciales. Cronquist’s Subclass Magnoliidae; Illiciales. APG III peripheral angiosperms; Superorder Austrobaileyanae. APG IV Order Austrobaileyales.

Species 47. Genera 2; Kadsura, Schisandra.

General remarks. For a monograph of Kadsura, see Saunders (1998).

Illustrations. • Le Maout and Decaisne: Schisandra glabra, as S. coccinea. • Schisandra propinqua: Hook. Ic. Pl. 18 (1887). • Schisandra propinqua: as Sphaerostema, Bot. Reg. 1688, 1835. Schisandra proquinqua. The stamens (fig. 2) nestle in small excavations in a solid globular mass (fig. 1), only the anthers being free.

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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