Family Menispermaceae

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; 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; without a persistent basal meristem. Domatia occurring in the family (5 genera); manifested as pits, or pockets, or hair tufts (rarely).

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

Leaf anatomy. Hydathodes present (occasionally), or absent.

The mesophyll with sclerencymatous idioblasts. Minor leaf veins without phloem transfer cells (Anamirta).

Stem anatomy. Cork cambium present; initially deep-seated, or superficial. Nodes tri-lacunar. Internal phloem absent. Secondary thickening usually anomalous; mostly via concentric cambia. ‘Included’ phloem mostly present. Xylem with tracheids; with vessels. Vessel end-walls simple. Wood parenchyma apotracheal (diffuse and in short tangential lines, and ‘conjunctive’ between the successive layers of bundles). Sieve-tube plastids S-type.

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); in cymes, in panicles, and in fascicles. The ultimate inflorescence unit 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 (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, biochemistry. Cyanogenic, or not cyanogenic. Alkaloids present (nearly always), or absent. Berberine present (in at least three genera). Iridoids not detected. Proanthocyanidins absent. Flavonols present, or absent; kaempferol. Ellagic acid absent (3 species, 3 genera). Aluminium accumulation not found. C3. C3 physiology recorded directly in Menispermum.

Geography, cytology. Temperate (warm), or 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 3 core angiosperms; peripheral eudicot; Superorder Ranunculanae; 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.
• Technical details (Cocculus, Coscinium, Menispermum, Stephania).
• Technical details: Cocculus (Thonner).
• Habit and technical details: Cissampelos (Lindley).
• Technical details: Cocculus (Lindley).

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