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The families of flowering plants

L. Watson and M.J. Dallwitz

Capparidaceae Juss.

Alternatively Capparaceae; ~ Cruciferae.

Including Cleomaceae Horan. (p.p.); excluding Cleomaceae, Koeberliniaceae, Pentadiplandraceae, Physenaceae, Setchellanthaceae.

Habit and leaf form. Trees, or shrubs, or lianas, or herbs (rarely); non-glandular, not resinous. ‘Normal’ plants. Self supporting, or climbing. Xerophytic (commonly), or mesophytic. Leaves alternate; spiral, or distichous (rarely); petiolate; non-sheathing; gland-dotted, or not gland-dotted; simple, or compound; when compound, palmate. Lamina when simple dissected, or entire; when simple/dissected, palmatifid. Leaves stipulate, or exstipulate (stipules small). Stipules (when present) often spiny (or represented by glands). Lamina margins often involute. Leaf development not ‘graminaceous’.

Leaf anatomy. The leaf lamina dorsiventral to centric. Abaxial epidermis papillose, or not papillose. Stomata present; anomocytic. Hairs of numerous kinds present (see illustration); mostly eglandular (by contrast with Cleomaceae); unicellular and multicellular. Complex hairs present, or absent; peltate, or stellate, or capitate. Adaxial hypodermis present, or absent. The mesophyll with sclerenchymatous idioblasts, or without sclerenchymatous idioblasts. Minor leaf veins without phloem transfer cells (Capparis, Steriphoma).

Axial (stem, wood) anatomy. Cork cambium present; initially deep-seated, or initially superficial. Nodes unilacunar. Internal phloem absent. Cortical bundles (secondary, pericyclic) present, or absent. Medullary bundles absent. Secondary thickening developing from a conventional cambial ring, or anomalous. The anomalous secondary thickening via concentric cambia (often), or from a single cambial ring.

The vessel end-walls simple. The vessels with vestured pits. The axial xylem with fibre tracheids, or without fibre tracheids; with libriform fibres; including septate fibres (rarely), or without septate fibres. The parenchyma paratracheal. The secondary phloem stratified into hard (fibrous) and soft (parenchymatous) zones, or not stratified. ‘Included’ phloem present (often), or absent. The wood partially storied, or not storied.

Reproductive type, pollination. Plants hermaphrodite, or dioecious. Pollination entomophilous.

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’; usually in racemes. The ultimate inflorescence units racemose. Inflorescences usually in racemes. Flowers bracteate; ebracteolate; regular to somewhat irregular; usually 4 merous. Floral receptacle developing a gynophore, or developing an androphore and developing a gynophore (causing the pistil to project). Hypogynous disk present; of separate members (a ring).

Perianth with distinct calyx and corolla; (4–)6 (rarely), or 8; 2–4 whorled (more or less resolvable into K2+2 and C4 with decussate calyx and diagonal petals, cf. Cruciferae). Calyx 4 (2+2); 2 whorled; basally gamosepalous, or polysepalous. Calyx lobes about the same length as the tube, or markedly longer than the tube. Calyx bilabiate, or regular; imbricate. Corolla 4 (diagonal, very rarely 2); 1 whorled; polypetalous (the petals equal or unequal, sometimes hooding).

Androecium basically 4, or 6–100 (i.e. to ‘many’, when branched). Androecial members branched (commonly), or unbranched; when branched, maturing centrifugally; free of the perianth; all equal to markedly unequal; basally coherent, or free of one another. The androecial bundles when detectable alternating with the corolla members. Androecium exclusively of fertile stamens, or including staminodes (or with staminodal branches). Stamens 4, or 6–20(–100) (to ‘many’); not didynamous, not tetradynamous; isomerous with the perianth to diplostemonous to polystemonous; basically oppositisepalous; filantherous, or petaloid and filantherous. Anthers dorsifixed to basifixed; dehiscing via longitudinal slits; introrse; tetrasporangiate. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or decussate. Anther wall initially with more than one middle layer. Tapetum glandular. Pollen grains aperturate; (2–)3(–4) aperturate; colporate (colporoidate); 2-celled.

Gynoecium 2 carpelled, or 10–12 carpelled (by dédoublement). The pistil 1 celled, or 2 celled, or 3–12 celled. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 1 locular (usually), or 2 locular (by false septa). Locules secondarily divided by ‘false septa’, or without ‘false septa’. Gynoecium when G=2 transverse. Ovary generally stipitate. Gynoecium non-stylate, or stylate. Styles 1; attenuate from the ovary; apical; shorter than the ovary. Stigmas 1; dorsal to the carpels, or commissural (?); dry type; papillate, or non-papillate; Group II type. Placentation when unilocular, parietal; when bilocular, parietal (without a ‘replum’, by contrast with Cleomaceae and Cruciferae). Ovules in the single cavity 10–100 (‘many’); when bilocular, 30–50 per locule (‘many’); arillate, or non-arillate; anatropous to campylotropous; bitegmic; crassinucellate. Outer integument contributing to the micropyle, or not contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral (usually small, large in Maerua). Synergids hooked (sometimes with filiform apparatus). Endosperm formation nuclear. Embryogeny onagrad, or solanad.

Fruit fleshy, or non-fleshy; dehiscent, or indehiscent; a capsule (often), or a berry (usually), or a drupe, or a nut, or a samara. Capsules sometimes valvular (resulting in two valves to which the placentas remain attached — i.e., no replum). Seeds endospermic, or non-endospermic. Cotyledons 2 (oily). Embryo variously curved, or bent. The radicle lateral, or dorsal.

Seedling. Germination phanerocotylar.

Physiology, phytochemistry. C3. C3 physiology recorded directly in Cadaba, Capparis, Maerua. Anatomy non-C4 type (Boscia, Capparis). Mustard-oils present. Cyanogenic, or not cyanogenic. Alkaloids present (21 species). Iridoids not detected. Proanthocyanidins absent. Flavonols absent. Ellagic acid absent (Euadenia). Aluminium accumulation not found. Sieve-tube plastids P-type, or S-type; when P-type type I (b).

Geography, cytology. Temperate to tropical. Widespread. X = 10, 11, 17.

Taxonomy. Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Violiflorae; Capparales. Cronquist’s Subclass Dilleniidae; Capparales. APG III core angiosperms; core eudicot; Superorder Rosanae; malvid. APG IV Order Brassicales.

Species 650. Genera about 26; Apophyllum, Atamisquea, Bachmannia, Belencita, Borthwickia, Boscia, Buchholzia, Cadaba, Capparis, Cladostemon, Crateva, Dactylaena, Dipterygium (or Cruciferae?), Euadenia, Forchhammeria, Maerua, Morisonia, Neothorelia, Podandrogyne, Poilanedora(?), Puccionia (or Cruciferae?), Ritchiea, Steriphoma, Stixis, Thilachium, Tirania.

General remarks. Separation from Cleomaceae may be unsustainable, since difficulties are encountered in assigning the genera. Precise comparative data on gynoecium and fruit structure are elusive or non-existent.

Economic uses, etc. Flower buds of Capparis spp. constitute culinary capers.

Illustrations. • Le Maout and Decaisne: Capparis. • Maerua angolensis, and Cynophalla flexuosa as Capparis sinclairii: Lindley. • Bachmannia woodii (as Niebuhria): Hook. Ic. Pl. 14 (1880–82). • Capparis acuminata: Bot. Reg. 1320, 1830. • Capparis brevispina: Hook. Ic. Pl. 2 (1837). • Capparis parvifolia, as C. murrayana: R. Wight, Ic. Pl. Indiae Orientalis 2 (1843). • Capparis rotundifolia, as C. pedunculosa: Hook. Ic. Pl. 2 (1837). • Cadaba linearifolia, as C. heterotricha: Hook. Ic. Pl. 9 (1852). • Maerua linearis, as Niebuhria: Hook. Ic. Pl. 2 (1837). • Atamisquea, Capparis, Cadaba, Steriphoma: leaf hairs (Solereder, 1908). • Capparis spp.: leaf anatomcal details (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.’.