The families of flowering plants.                                                                                                                                                                

Cruciferae Juss.


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AlternativelyBrassicaceae Burnett (nom. altern.)

IncludingRaphanaceae Horan., Stanleae (Stanleyaceae) Nutt.

Habit and leaf form. Herbs (mostly), or shrubs (rarely), or lianas (rarely). ‘Normal’ plants (mostly), or switch-plants (e.g. Caulanthus, sometimes spiny, e.g. Vella). Plants non-succulent (mainly), or succulent (a few, e.g. Cakile). Annual, biennial, and perennial; with a basal aggregation of leaves, or with neither basal nor terminal aggregations of leaves. Self supporting (usually), or climbing (rarely). When shrubby, often pachycaul. Hydrophytic, helophytic, mesophytic, and xerophytic; when hydrophytic, rooted. Leaves of hydrophytes submerged and emergent. Heterophyllous, or not heterophyllous. Leaves minute to very large; alternate; spiral (usually), or distichous (rarely); fleshy (occasionally), or ‘herbaceous’; petiolate, or subsessile, or sessile; sheathing to non-sheathing; foetid (sometimes), or without marked odour; simple (usually), or compound (rarely); epulvinate; when compound, pinnate (with articulated leaflets). Lamina when simple, dissected to entire; when simple-dissected, pinnatifid, or runcinate; one-veined (rarely), or pinnately veined; cross-venulate. Leaves exstipulate. Lamina margins entire, or serrate, or dentate. Leaves without a persistent basal meristem.

Leaf anatomy. Mucilaginous epidermis present (commonly), or absent.

Lamina dorsiventral, or isobilateral, or centric. Minor leaf veins without phloem transfer cells (4 genera).

Stem anatomy. Cork cambium present, or absent; initially in woody species deep-seated. Nodes unilacunar, or tri-lacunar, or multilacunar. Cortical bundles present (rarely), or absent. Medullary bundles present (rarely), or absent. Secondary thickening absent, or developing from a conventional cambial ring, or anomalous (?— given the occasional presence of cortical and/or medullary bundles); usually from a single cambial ring. ‘Included’ phloem present (e.g.Brassica, Cochlearia), or absent. Xylem with fibre tracheids; with vessels. Vessel end-walls simple. Vessels with vestured pits. Wood partially storied, or not storied (?); parenchyma paratracheal (scarce). Sieve-tube plastids P-type, or S-type; when P-type type I (b).

Reproductive type, pollination. Unisexual flowersabsent. Plants hermaphrodite. Floral nectaries present (nearly always), or absent. Nectar secretion from the perianth, from the disk, and from the androecium. Pollination anemophilous (e.g.Pringlea), or entomophilous (mostly).

Inflorescence, floral, fruit and seed morphology. Flowers aggregated in ‘inflorescences’ (usually), or solitary (rarely); in corymbs (commonly), or in racemes, or in spikes, or in fascicles. The ultimate inflorescence unit racemose. Inflorescences terminal, or axillary, or leaf-opposed (e.g. Coronopus); sometimes more or less pseudanthial, or not pseudanthial. Flowers ebracteate (usually), or bracteate; ebracteolate (mostly), or bracteolate; minute to medium-sized; fragrant, or odourless; more or less regular (usually), or somewhat irregular. The floral irregularity (when noticeable) involving the perianth (the outer petals of pseudanthia sometimes radiate and larger). Flowers 2 merous; cyclic; polycyclic (usually 6), or tetracyclic to pentacyclic (when C or A whorls lacking). Floral receptaclenearly always with neither androphore nor gynophore (by contrast with Capparidaceae — the long, slender gynophore of Stanleyeae being exceptional). Free hypanthium absent. Hypogynous disk present (nearly always), or absent; extrastaminal, or intrastaminal, or extrastaminal and intrastaminal (usually between and/or external to the filaments, sometimes surrounding their bases); of separate members (usually, in variable configurations which are of taxonomic interest), or annular (in that the members are sometimes confluent).

Perianthwith distinct calyx and corolla, or sepaline (the corolla occasionally lacking); 8 (usually), or 4; characteristically 3 whorled (K 2+2 decussate with the outer pair median, C4, the corolla diagonal), or 2 whorled (when corolla lacking). Calyx 2 whorled (generally, supposedly — usually obviously so in bud); polysepalous; regular; members sometimes dimorphic, with the lateral (inner) pair often basally saccate for nectar storage; decussate. Corolla 4 (diagonal); 1 whorled; polypetalous; imbricate, or contorted; regular (usually); white, or yellow, or orange, or purple, or blue. Petals clawed (usually), or sessile (e.g. Cuphonotus, Geococcus).

Androecium 6 (usually), or 2–4, or 8–16 (rarely). Androecial members branched (usually, in that the inner whorl of 4 is derived from only 2 primordia), or unbranched; free of the perianth; markedly unequal (usually, the outer pair shorter), or all equal; free of one another, or coherent (the inner four sometimes basally connate in pairs); 2 whorled (usually, 2+4), or 1 whorled. Androecium exclusively of fertile stamens. Stamens 6 (usually), or 2–4, or 8–16 (rarely); tetradynamous (nearly always), or not didynamous, not tetradynamous (when the outer pair missing); oppositisepalous. Filaments appendiculate, or not appendiculate. Anthers basifixed; non-versatile; dehiscing via longitudinal slits; introrse; unilocular to bilocular; tetrasporangiate; variously appendaged, or unappendaged. Endothecium developing fibrous thickenings. Anther epidermis persistent. Microsporogenesis simultaneous. The initial microspore tetrads tetrahedral, or isobilateral, or decussate. Anther wall initially with one middle layer, or initially with more than one middle layer; of the ‘monocot’ type. Tapetum glandular. Pollen shed as single grains. Pollen grains aperturate, or nonaperturate (occasionally); 3(–10) aperturate; colpate; 3-celled (in 16 genera).

Gynoecium2 carpelled. The pistil 1–2 celled. Gynoecium syncarpous; eu-syncarpous; superior. Ovary 1–2 locular. Locules secondarily divided by ‘false septa’ (usually, as evidenced by the longitudinal splitting of the parietal placentas by the replum), or without ‘false septa’ (e.g. Menkea). Gynoecium transverse. Ovary sessile (usually), or stipitate (rarely, cf. Capparidaceae). Gynoecium non-stylate, or non-stylate to stylate. Styles 1 (more or less reduced); apical. Stigmas 1–2; commissural (mostly, when detectably two or bilobed), or dorsal to the carpels (e.g. Matthiola, Moricandia); 1 lobed, or 2 lobed; more or less capitate; dry type; papillate; Group II type.Placentation when unilocular, parietal; when bilocular, parietal (but peculiarly so, the usual two placentas usually each longitudinally divided by a ‘false’ septum, the ‘replum’, separating the locules). Ovules in the single cavity 1–50 (to ‘many’); (1–)3–50 per locule (i.e. usually several to ‘many’ per placenta); commonly pendulous, or horizontal; with ventral raphe; non-arillate; anatropous; bitegmic; tenuinucellate, or crassinucellate. Outer integument contributing to the micropyle. Embryo-sac development Polygonum-type. Polar nuclei fusing prior to fertilization. Antipodal cells formed; 3; not proliferating; ephemeral. Synergids hooked (sometimes with filiform apparatus). Endosperm formation nuclear. Embryogeny onagrad.

Fruit non-fleshy; dehiscent (usually), or indehiscent (Cakile, Crambe, Coronopus etc), or a schizocarp (e.g. sometimes in Coronopus), or lomentaceous (or sometimes partially so). Mericarps when schizocarpic, 2. Fruit a silicula to a siliqua (with variations on these themes, sometimes partially of entirely lomentaceous), or capsular-indehiscent (i.e. when neither dehiscent nor lomentaceous). Capsules when dehiscent, valvular (the valves usually falling or spreading upwards to reveal the replum). Seeds scantily endospermic, or non-endospermic; small to medium sized; winged (e.g. Matthiola, Isatis), or wingless. Embryo well differentiated. Cotyledons 2; flat, or folded (longitudinally or transversely). Embryo chlorophyllous (32/52); bent (nearly always folded, cotyledons against radicle), or straight (e.g. Leavenworthia). The radicle lateral, or dorsal.

Seedling.Germination phanerocotylar, or cryptocotylar.

Physiology, biochemistry. Mustard-oilspresent. Cyanogenic, or not cyanogenic. Alkaloids present, or absent (more often). Iridoids not detected. Proanthocyanidins absent (except sometimes in the seedcoat). Flavonols present, or absent; when present, kaempferol and quercetin. Ellagic acid absent (9 species, 7 genera). Arbutin absent. Aluminium accumulation not found. C3 (almost exclusively), or C3-C4 intermediate. C3 physiology recorded directly in Barbarea, Berteroa, Brassica,Cakile, Capsella, Cheiranthus, Crambe, Dipterygium,Farsetia, Matthiola, Raphanus, Sisymbrium, Thlaspi. C3-C4 intermediacy in Moricandia (3 species). Anatomy non-C4 type (Aethionema, Alyssum, Arabis, Brassica, Capsella,Cardamine, Descurainia, Diceratella, Draba, Erysimum,Erucastrum, Farsetia, Fortuynia, Leavenworthia, Lepidium,Lesquerella, Matthiola, Morieria, Oreophyton, Rorippa,Sisymbrium, Thlaspi etc.).

Geography, cytology. Frigid zone to tropical. Cosmopolitan, concentrated in the North temperate and Mediterranean. X = 5–12(+).

Taxonomy.Subclass Dicotyledonae; Crassinucelli. Dahlgren’s Superorder Violiflorae; Capparales. Cronquist’s Subclass Dilleniidae; Capparales. APG 3 core angiosperms; core eudicot; Superorder Rosanae; malvid; Order Brassicales.

Species 3200. Genera about 350; Acanthocardamum, Aethionema, Agallis,Alliaria, Alyssoides, Alysopsis, Alyssum, Ammosperma,Anastatica, Anchonium, Andrzeiowskia, Anelsonia, Aphragmus,Aplanodes, Arabidella, Arabidopsis, Arabis, Arcyosperma,Armoracia, Aschersoniodoxa, Asperuginoides, Asta, Atelanthera,Athysanus, Aubretia, Aurinia, Ballantinia, Barbarea,Berteroa, Berteroella, Biscutella, Bivonaea, Blennodia,Boleum, Boreava, Bornmuellera, Borodinia, Botscantzevia,Brachycarpaea, Brassica, Braya, Brayopsis, Brossardia,Bunias, Cakile, Calepina, Calymmatium, Camelina,Camelinopsis, Capsella, Cardamine, Cardaminopsis, Cardaria,Carinavalva, Carrichtera, Catadysia, Catenulina, Caulanthus,Caulostramina, Ceratocnemum, Ceriosperma, Chalcanthus,Chamira, Chartoloma, Cheesemania, Cheiranthus (= Erisymum),Chlorocrambe, Chorispora, Christolea, Chrysobraya, Chrysochamela,Cithareloma, Clastopus, Clausia, Clypeola, Cochlearia,Coelonema, Coincya, Coluteocarpus, Conringia, Cordylocarpus,Coronopus, Crambe, Crambella, Cremolobus, Cryptospora,Cuphonotus, Cusickiella, Cycloptychis, Cymatocarpus,Cyphocardamum, Dactylocardamum, Degenia, Delpinophytum,Descurainia, Diceratella, Dichasianthus, Dictyophragmus,Didesmus, Didymophysa, Dielsiocharis, Dilophia, Dimorphocarpa,Diplotaxis, Dipoma, Diptychocarpus, Dithyrea, Dolichirhynchus,Dontostemon, Douepea, Draba, Drabastrum, Drabopsis,Dryopetalon, Eigia, Elburzia, Enarthrocarpus, Englerocharis,Eremobium, Eremoblastus, Eremodraba, Eremophyton, Ermania,Ermaniopsis, Erophila, Erucaria, Erucastrum, Erysimum,Euclidium, Eudema, Eutrema, Euzomodendron, Farsetia,Fezia, Fibigia, Foleyola, Fortuynia, Galitzkya,Geococcus, Glaribraya, Glastaria, Glaucocarpum, Goldbachia,Gorodkovia, Graellsia, Grammosperma, Guiraoa, Gynophorea,Halimolobos, Harmsiodoxa, Hedinia, Heldreichia, Heliophila,Hemicrambe, Hemilophia, Hesperis, Heterodraba, Hirschfeldia,Hollermayera, Hornungia, Hornwoodia, Hugueninia, Hymenolobus,Iberis, Idahoa, Iodanthus, Ionopsidium, Irenepharsus,Isatis, Ischnocarpus, Iskandera, Iti, Ivania,Kernera, Kremeriella, Lachnocapsa, Lachnoloma, Leavenworthia,Lepidium, Lepidostemon, Leptaleum, Lesquerella, Lignariella,Lithodraba, Lobularia, Lonchophora, Loxostemon, Lunaria,Lyocarpus, Lyrocarpa, Macropodium, Malcolmia, Mancoa,Maresia, Mathewsia, Matthiola, Megacarpaea, Megadenia,Menkea, Menonvillea, Microlepidium, Microsysymbrium,Microstigma, Morettia, Moricandia, Moriera, Morisia,Murbeckiella, Muricaria, Myagrum, Nasturtiopsis, Nasturtium (= Rorippa), Neomartinella, Neotchihatchewia, Neotorularia,Nerisyrenia, Neslia, Neuontobotrys, Notoceras, Notothlaspi,Ochthodium, Octoceras, Onuris, Oreoloma, Oreophyton,Ornithocarpa, Orychophragmus, Otocarpus, Oudneya, Pachycladon,Pachymitus, Pachyphragma, Pachypterygium, Parlatoria,Parodiodoxa, Parolinia, Parrya, Parryodes, Pegaeophyton,Peltaria, Peltariopsis, Pennellia, Petiniotia, Petrocallis,Phaeonychium, Phlebolobium, Phlegmatospermum, Phoenicaulis,Physaria, Physocardamum, Physoptychis, Physorrhynchus,Platycraspedum, Polyctenium, Polypsecadium, Pringlea,Prionotrichon, Pritzelago, Pseuderucaria, Pseudocamelina,Pseudoclausia, Pseudofortuynia, Pseudovesicaria, Psychine,Pterygiosperma, Pterygostemon, Pugionium, Pycnoplinthopsis,Pycnoplinthus, Pyramidium, Quezeliantha, Quidproquo,Raffenaldia, Raphanorhyncha, Raphanus, Rapistrum, Reboudia,Redowskia, Rhizobotrya, Ricotia, Robeschia, Rollinsia,Romanschulzia, Roripella, Rorippa, Rytidocarpus, Sameraria,Sarcodraba, Savignya, Scambopus, Schimpera, Schivereckia,Schizopetalon, Schlechteria, Schoenocrambe, Schouwia,Scoliaxon, Selenia, Sibara, Silicularia, Sinapidendron,Sinapis, Sisymbrella, sisymbriopsis, Sisymbrium, Smelowskia,Sobolewslia, Sohms-Laubachia, Sophiopsis, Sphaerocardamum,Spirorhynchus, Spryginia, Staintoniella, Stanfordia,Stanleya, Stenopetalum, Sterigmostemum, Stevenia, Straussiella,Streptanthella, Streptanthus, Streptoloma, Stroganowia,Stubebdorffia, Subularia, Succowia, Synstemon, Synthlipsis,Taphrospermum, Tauscheria, Teesdalia, Teesdaliopsis,Tetracme, Thelypodiopsis, Thelypodium, Thlaspeocarpa,Thlaspi, Thysanocarpus, Trachystoma, Trichotolinum,Trochiscus, Tropidocarpum, Turritis, Vella, Warea,Weberbauera, Werdermannia, Winklera, Xerodraba, Yinshania,Zerdana, Zilla.

General remarks. For family review, see Vaughan, Macleod and Jones 1976.

Economic uses, etc. Important food crops from Brassica (cabbage, cauliflower, broccoli, kohlrabi, turnips, Brussels sprouts); Raphanus (radish); Rorippa (watercress); condiments from Brassica (mustard), Armoracia (horse-radish); ornamentals from about 50 genera, e.g. Arabis, Erysimum (Cheiranthus), Hesperis, Iberis, Lobularia, Lunaria,Matthiola.


  • Technical details: Cheiranthus, Cochlearia, Parolinia.
  • Technical details: Arabis, Barbarea, Lunaria, Tetrapoma (~Rorippa).
  • Technical details: Cremolobus, Erophila, Eruca, Megacarpaea, Vesicaria (~Alyssoides).
  •   Technical details: Aethionema, Iberis, Isatis, Capsella, Lepidium, Neslia, Myagrum, Thlaspi.
  • Technical details: Bunias, Clypeola, Crambe, Myagrum, Raphanistrum (~Raphanus), Thysanocarpus.
  •  Arabidopsis thaliana (as Draba thaliana): Eng. Bot. 115, 1863.
  •  Brassica rapa (Turnip) and Brassica rapa ssp. campestris (Wild Turnip) : Eng. Bot. 89 and 90, 1863.
  • Brassica oleracea (Wild Cabbage) and Brassica napus (Rape): Eng. Bot. 87 and 88, 1863.
  •  Cakile maritima and Crambe maritima: Eng. Bot. 134, 1863.
  • Cardamine amara, C. pratensis and C. hirsuta: Eng. Bot. 108–110, 1863.
  •  Erophila verna (as 3 forms of Draba verna): Eng. Bot. 134, 1863.
  •  Isatis tinctoria: B. Ent. 734.
  •  Rorippa nasturtium-aquatica (Watercress, as Nasturtium officinalis): Eng. Bot. 125, 1863.
  •  Brassiceae (Brassica, Diplotaxis: B. Ent. compilation).
  • Brassiceae (Rhynchosinapis, Cakile, Sinapis: B. Ent. compilation).  
  • Lepidieae (Capsella, Coronopus, Cochlearia: B. Ent. compilation).
  •  Lepidieae (Teesdalia), Drabeae (Erophila), Matthioleae (Matthiola): B. Ent. compilation.  
  • Arabideae (Arabis, Cardamine, Barbarea: B. Ent. compilation).
  • Arabideae (Rorippa, Dentaria: B. Ent. compilation).
  • Hesperideae (Erysimum, Cheiranthus, Hesperis: B. Ent. compilation).
  • Sisymbrieae (Arabidopsis, Descurainia, Alliaria: B. Ent. compilation).
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