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The families of flowering plants

L. Watson and M.J. Dallwitz

Menispermaceae Juss.

Habit and leaf form. Scandent shrubs, or lianas, or herbs (rarely), or trees (rarely). Climbing (usually), or self supporting; mostly stem twiners; Stephania twining clockwise. Mesophytic. Leaves alternate; spiral (usually with serial axillary buds); petiolate; non-sheathing; not gland-dotted; simple (usually), or compound (rarely); peltate (sometimes), or not peltate; when compound, ternate. Lamina when simple, entire (usually), or dissected; when simple/dissected, palmatifid; pinnately veined, or palmately veined; cross-venulate. Leaves exstipulate; leaf development not ‘graminaceous’. Domatia occurring in the family (known from 5 genera); manifested as pits, or pockets, or hair tufts (rarely).

General anatomy. Plants with ‘crystal sand’, or without ‘crystal sand’.

Leaf anatomy. The leaf lamina dorsiventral (usually), or centric (rarely). Hydathodes present (occasionally), or absent. Hairs present; eglandular and glandular (the former uniseriate, of two or more cells, the latter variously small, unicellular and ellipsoidal or clavate, or long uniseriate, or shaggy). Complex hairs absent. The mesophyll with sclerenchymatous idioblasts. Minor leaf veins without phloem transfer cells (Anamirta).

Axial (stem, wood) anatomy. Young stems with solid internodes. Pith with diaphragms (sometimes with thin transverse septa), or without diaphragms; homogeneous to heterogeneous. Cork cambium present; initially deep-seated, or initially superficial. Nodes tri-lacunar. Primary vascular tissues comprising a ring of bundles (the bundles in young stems distinct in stem sections even to the unaided eye); collateral. Internal phloem absent. Cortical bundles absent. Medullary bundles absent. Secondary thickening anomalous (usually), or developing from a conventional cambial ring. The anomalous secondary thickening when manifest, mostly via concentric cambia (but often eccentric, see illustration). Primary medullary rays wide. The axial xylem with vessels.

The wood diffuse porous. The vessels small to medium; solitary, radially paired, and in radial multiples (but mostly solitary). The vessel end-walls horizontal to oblique (slightly oblique); simple. The vessels without spiral thickening. The axial xylem with tracheids; without septate fibres. The fibres without spiral thickening. The parenchyma apotracheal (diffuse and in short tangential lines, and ‘conjunctive’ between the successive layers of bundles). ‘Included’ phloem mostly present. The wood partially storied.

Reproductive type, pollination. Fertile flowers functionally male, or functionally female, or functionally male and functionally female. Plants dioecious (mostly), or monoecious (?). Female flowers with staminodes, or without staminodes.

Inflorescence, floral, fruit and seed morphology. Flowers solitary (rarely), or aggregated in ‘inflorescences’ (sometimes cauliflorous); usually in cymes, in panicles, and in fascicles. The ultimate inflorescence units cymose. Inflorescences axillary, or cauliflorous. Flowers small (rarely brightly coloured); regular (usually), or somewhat irregular; more or less 3 merous; cyclic; often approximating to K3+3, C3+3, A3+3, G3 — but with many exceptions. Floral receptacle developing a gynophore (sometimes), or with neither androphore nor gynophore. Free hypanthium absent.

Perianth with distinct calyx and corolla (usually), or sepaline (occasionally without petals); (1–)6(–12), or (4–)12(–24); 2–6 whorled; isomerous, or anisomerous. Calyx (1–)6(–12); often 2 whorled; usually polysepalous; imbricate, or valvate. Corolla when present (i.e. usually), (3–)6(–12); often 2 whorled; polypetalous (usually), or gamopetalous (rarely).

Androecium (1–)6(–40). Androecial members branched, or unbranched; free of the perianth; free of one another, or coherent (the filaments often connate or in bundles); when joined 1 adelphous, or 2–5 adelphous (?); often 2 whorled. Androecium exclusively of fertile stamens. Stamens (1–)6(–40); reduced in number relative to the adjacent perianth to isomerous with the perianth to polystemonous; alternisepalous (often), or oppositisepalous. Anthers dehiscing via longitudinal slits, or dehiscing transversely; introrse; 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 (two); of the ‘dicot’ type. Tapetum glandular. Pollen grains aperturate (usually), or nonaperturate; (2–)3(–4) aperturate; colpate, or colporate, or rugate (rug(or)ate); 2-celled (in Cocculus and Hyperbaena).

Gynoecium (1–)3 carpelled, or 6–30 carpelled (in one or more whorls). The pistil when monomerous, 1 celled. Gynoecium monomerous, or apocarpous; of one carpel, or eu-apocarpous; superior. Carpel apically stigmatic; 2 ovuled (with one abortive). Placentation marginal (ventral). Stigmas dry type; non-papillate; Group II type. Ovules pendulous to horizontal; anatropous to amphitropous, or hemianatropous; unitegmic, or bitegmic; crassinucellate. Outer integument not contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; proliferating (Tinospora), or not proliferating; ephemeral, or persistent. Synergids pear-shaped, or hooked (sometimes with filiform apparatus). Endosperm formation nuclear. Embryogeny onagrad.

Fruit fleshy to non-fleshy; an aggregate (usually, in a head), or not an aggregate (where only one of the carpels develops). The fruiting carpels not coalescing (each usually curved). The fruiting carpel indehiscent; nucular, or drupaceous (usually curved, often horseshoe shaped). Fruit 1 seeded. Seeds endospermic, or non-endospermic. Endosperm when present, ruminate, or not ruminate; oily. Cotyledons 2. Embryo straight to curved.

Seedling. Germination phanerocotylar, or cryptocotylar.

Physiology, phytochemistry. C3. C3 physiology recorded directly in Menispermum. Cyanogenic, or not cyanogenic. Alkaloids present (nearly always), or absent. Berberine present (in at least three genera, including Tinospora). Iridoids not detected. Proanthocyanidins absent. Flavonols present, or absent; kaempferol. Ellagic acid absent (3 species, 3 genera). Aluminium accumulation not found. Sieve-tube plastids S-type.

Geography, cytology. Temperate (warm), sub-tropical to tropical. Pantropical and warm. X = 11–13, 19, 25.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Ranunculiflorae; Ranunculales. Cronquist’s Subclass Magnoliidae; Ranunculales. APG III core angiosperms; peripheral eudicot; Superorder Ranunculanae. APG IV Order Ranunculales.

Species 520. Genera about 75; Abuta, Albertisia, Anamirta, Anisocycla, Anomospermum, Antizoma, Arcangelisia, Aspidocarya, Beirnaertia, Borismene, Burasaia, Calycocarpum, Carronia, Caryomene, Chasmanthera, Chlaenandra, Chondrodendron, Cionomene, Cissampelos, Cocculus, Coscinium, Curarea, Cyclea, Dialytheca, Dioscoreophyllum, Diploclisia, Disciphania, Elephantomene, Eleutharrhena, Fibraurea, Haematocarpus, Hyperbaena, Hypserpa, Jateorhiza, Kolobopetalum, Legnephora, Leptoterantha, Limacia, Limaciopsis, Macrococculus, Menispermum, Odontocarya, Orthogynium, Orthomene, Pachygone, Parabaena, Penianthus, Pericampylus, Platytinospora, Pleogyne, Pycnarrhena, Rhaptonema, Rhigiocarya, Sarcolophium, Sarcopetalum, Sciadotenia, Sinomenium, Sphenocentrum, Spirospermum, Stephania, Strychnopsis, Synandropus, Synclisia, Syntriandrum, Syrrheonema, Telitoxicum, Tiliacora, Tinomiscium, Tinospora, Triclisia, Ungulipetalum.

Illustrations. • Le Maout and Decaisne: Cocculus, Coscinium, Menispermum, Stephania. • Cocculus leaeba: Thonner. • Cissampelos tropaeofolia: Lindley. • Cocculus macrocarpus: Lindley. • Diploclisia glaucescens (as Cocculus macrocarpus): R. Wight (1840). • Hypserpa nitida: Trimen, Ill. Fl Ceylon (1893). • Tinospopra sagittata, as Limacia: Hook. Ic. Pl. 18 (1888). • Anomospermum grandifolium: TS stem, with anomalous, asymmetric secondary thickening (Solereder, 1908).

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: 15th April 2018.’.