| # The Families of Angiosperms |

L. Watson and M.J. Dallwitz

### Leeaceae (DC.) Dum.

~ Vitaceae-Leeoideae.

Habit and leaf form. Herbs, or trees, or shrubs (the branches sometimes prickly). Self supporting (without tendrils, by contrast with Vitaceae s. str.). Leaves alternate (usually), or opposite; petiolate; non-sheathing; simple (rarely), or compound; unifoliolate (rarely), or ternate (rarely), or pinnate, or bipinnate to multiply compound (to tripinnate); when pinnate, imparipinnate. Lamina pinnately veined; cross-venulate. Leaves exstipulate (but the petioles sometimes exhibiting auricles or sheathing expansions near the base). Lamina margins usually dentate.

Leaf anatomy. The leaf lamina dorsiventral. Leaves with ‘pearl glands’ (commonly, these deciduous: see illustration), or without ‘pearl glands’. Stomata present; anomocytic (rarely), or cyclocytic, or actinocytic. The mesophyll containing mucilage cells (with or without raphides); containing crystals. The crystals raphides and druses. Minor leaf veins without phloem transfer cells.

Axial (stem, wood) anatomy. Secondary thickening developing from a conventional cambial ring. Primary medullary rays wide.

The vessel end-walls simple. The parenchyma paratracheal.

Reproductive type, pollination. Unisexual flowers absent. Plants hermaphrodite.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’. The ultimate inflorescence units cymose. Inflorescences terminal (usually, usually erect), or axillary (rarely); usually large, corymbose and many-flowered, often rusty-tomentose. Flowers small; regular; (3–)5 merous; cyclic; tetracyclic. Free hypanthium absent.

Perianth with distinct calyx and corolla; (8–)10; 2 whorled; isomerous. Calyx (4–)5; 1 whorled; gamosepalous; shortly toothed; cupuliform; regular; valvate. Corolla (4–)5; 1 whorled; gamopetalous; valvate; regular (the petals often reflexed at anthesis).

Androecium (4–)5. Androecial members basally adnate (to the corolla); all equal; coherent (the filaments connate above the fusion with the corolla, forming a tube which sometimes has entire or bifid lobes alternating with the anthers, and which sometimes proliferates internally near the middle to form a pendulous, tubular-obconic membrane); 1 adelphous; 1 whorled. Androecium exclusively of fertile stamens. Stamens (4–)5; isomerous with the perianth; alternisepalous; opposite the corolla members. Anthers introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral. Anther wall initially with more than one middle layer (2). Tapetum glandular. Pollen grains aperturate; 3 aperturate; colporate; psilate; 2-celled.

Gynoecium 2–3(–4) carpelled (with each primary ‘carpel’ incompletely divided lengthwise by a ‘false septum’ from its midrib), or 4–6(–8) carpelled (if the same structure be so interpreted). Carpels reduced in number relative to the perianth, or isomerous with the perianth, or increased in number relative to the perianth (depending on the interpretation). The pistil 2–8 celled. Gynoecium syncarpous; eu-syncarpous; superior (but often somewhat embedded in a disk). Ovary 4–6(–8) locular. Locules secondarily divided by ‘false septa’, or without ‘false septa’ (depending on interpretation). Gynoecium stylate. Styles 1. Stigmas 1; capitate; dry type; papillate; Group II type. Placentation basal. Ovules 1 per locule (i.e. in each chamber, though the ‘carpels’ may be interpreted as biovulate); sunken in the placenta (often); ascending; anatropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Endosperm formation nuclear. Embryogeny asterad.

Fruit fleshy; indehiscent; a berry (with sometimes thin flesh). Seeds endospermic. Endosperm ruminate; oily. Cotyledons 2.

Seedling. Germination phanerocotylar.

Physiology, phytochemistry. Sugars transported as sucrose. Not cyanogenic. Iridoids not detected. Saponins/sapogenins absent. Proanthocyanidins present; cyanidin and delphinidin. Flavonols present; kaempferol, quercetin, and myricetin. Ellagic acid absent. Aluminium accumulation not found. Sieve-tube plastids P-type; type I (b).

Geography, cytology. Paleotropical. Tropical. Palaeotropical.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Santaliflorae; Vitidales. Cronquist’s Subclass Rosidae; Rhamnales. APG III core angiosperms; cf Superorder Rosanae (?). APG IV Order Vitales (as a synonym of Vitaceae).

Species 70. Genera 1; only genus, Leea.

General remarks. The compiled data have Leea differing from Vitidaceae (q.v.) in the exstipulate leaves without gland-dots, the valvate calyx, the adnate and cohering stamens, the gynoecium with one basal ovule per locule, and the outer integument of the ovule contributing to the micropyle; also in records of papillate stigmas and sugars transported entirely as sucrose.

Illustrations. • Leea coccinea: Bot. Mag. 88 (1862). • Leea aequata: TS leaf and details of pearl gland (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: 8th June 2026. delta-intkey.com’.

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