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The Families of Angiosperms

L. Watson and M.J.Dallwitz

Cruciferae Juss.

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); mostly autotrophic (?), or utilizing the nutritious exudates oftrapped or harboured animals (in that mucilage secreted byCapsellabursa-pastoris seeds contains compounds that attract soil organisms, toxinswhich kill them, and digestive enzymes). Trapping mechanism passive. With abasal aggregation of leaves, or without conspicuous aggregations of leaves, orwith terminal aggregations of leaves (rarely). Self supporting (usually), orclimbing (rarely). When shrubby, often pachycaul. Hydrophytic, helophytic,mesophytic, and xerophytic; when hydrophytic, rooted. Leaves of hydrophytessubmerged and emergent. Conspicuously heterophyllous, or not conspicuouslyheterophyllous.Leaves minute to very large; alternate, or opposite towhorled (rarely); 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; whensimple-dissected, pinnatifid, or runcinate; one-veined (rarely), or pinnatelyveined; cross-venulate. Leaves exstipulate. Lamina margins entire, or serrate,or dentate. Leaf development not ‘graminaceous’.

Leaf anatomy.The leaflamina dorsiventral, or bifacial, or centric. Mucilaginous epidermis present(commonly), or absent. Hairs of numerous kinds present (in the family; mostlyunicellular but these variously simple, forked, Y- or T- shaped, sometimesstellate: see illustration); eglandular (mostly), or glandular (found in only afew genera, e.gHesperis,Matthiola). Minor leaf veins withoutphloem transfer cells (4 genera).

Axial (stem, wood) anatomy.Young stems with solid internodes, or with hollow internodes(commonly). Pith mostly without diaphragms; mostly homogeneous. Cork cambiumpresent (at least in woody forms), or absent; when present, initiallydeep-seated. Nodes unilacunar, or tri-lacunar, or multilacunar. Primary vasculartissues variously in a cylinder, without separate bundles (with a continuousring of cambium, and recognisable medullary rays much reduced or absent), orcomprising a ring of bundles (with or without subsequent interfascicular cambialactivity); collateral. Cortical bundles present (rarely), or absent. Medullarybundles present (rarely), or absent. Secondary thickening absent, or developingfrom a conventional cambial ring (the lignified interfascicular tissue usuallylacking vessels), or anomalous (occasionally). The anomalous secondarythickening when present via concentric cambia, or from a single cambial ring.

The vessels solitary, or solitary, in radial multiples, and clustered. Thevessel end-walls simple. The vessels with vestured pits. The axial xylem withfibre tracheids; without septate fibres. The fibres without spiral thickening.The parenchyma paratracheal (scarce). ‘Included’ phloem present(e.g.Brassica,Cochlearia), or absent. The wood partially storied,or not storied (?).

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

Inflorescence, floral, fruit and seedmorphology.Flowers aggregated in ‘inflorescences’(usually), or solitary (rarely); in corymbs (commonly), or in racemes, or inspikes, or in fascicles. The ultimate inflorescence units 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 tomedium-sized; fragrant, or odourless; more or less regular (usually), orsomewhat irregular. The floral irregularity (when noticeable) involving theperianth (the outer petals of pseudanthia sometimes radiate and larger). Flowers2 merous; cyclic; polycyclic (usually 6), or tetracyclic to pentacyclic (when Cor A whorls lacking).Floral receptaclenearly alwayswithneither androphore nor gynophore (by contrast withCapparidaceae —the long, slender gynophore of Stanleyeae being exceptional). Freehypanthium absent. Hypogynous disk present (nearly always), or absent;extrastaminal, or intrastaminal, or extrastaminal and intrastaminal (usuallybetween and/or external to the filaments, sometimes surrounding their bases); ofseparate members (usually, in variable configurations which are of taxonomicinterest), or annular (in that the members are sometimes confluent).

Perianth with distinct calyx and corolla, or sepaline (thecorolla occasionally lacking); 8 (usually), or 4;characteristically3whorled (K 2+2 decussate with the outer pair median, C4, the corolla diagonal),or 2 whorled (when corolla lacking).Calyx2 whorled (generally,supposedly — usually obviously so in bud); polysepalous; regular;members sometimes dimorphic, with the lateral (inner) pair often basally saccatefor nectar storage; decussate. Corolla 4 (diagonal); 1 whorled; polypetalous;imbricate, or contorted; regular (usually); white, or yellow, or orange, orpurple, 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 isderived from only 2 primordia), or unbranched; free of the perianth; markedlyunequal (usually, the outer pair shorter), or all equal; free of one another, orcoherent (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 theouter pair missing); oppositisepalous. Filaments appendiculate, or notappendiculate. Anthers basifixed; non-versatile; dehiscing via longitudinalslits; introrse; unilocular to bilocular; tetrasporangiate; variouslyappendaged, or unappendaged. Endothecium developing fibrous thickenings. Antherepidermis persistent. Microsporogenesis simultaneous. The initial microsporetetrads tetrahedral, or isobilateral, or decussate. Anther wall initially withone 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).

Gynoecium 2 carpelled. The pistil 1–2 celled.Gynoeciumsyncarpous;eu-syncarpous; superior. Ovary 1–2 locular. Loculessecondarily divided by ‘false septa’ (usually, as evidenced by thelongitudinal splitting of the parietal placentas by the replum), or without‘false septa’ (e.g.Menkea). Gynoecium transverse. Ovarysessile (usually), or stipitate (rarely, cf.Capparidaceae). Gynoeciumnon-stylate, or non-stylate to stylate. Styles 1 (more or less reduced); apical.Stigmas 1–2;commissural (mostly, when detectably two orbilobed), 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;whenbilocular,parietal (but peculiarly so, the usual two placentas usuallyeach longitudinally divided by a ‘false’ septum, the‘replum’, separating the locules). Ovules in the single cavity1–50 (to ‘many’); (1–)3–50 per locule (i.e. usuallyseveral to ‘many’ per placenta); commonly pendulous, or horizontal;with ventral raphe; non-arillate; anatropous; bitegmic; tenuinucellate, orcrassinucellate. Outer integument contributing to the micropyle. Embryo-sacdevelopmentPolygonum-type. Polar nuclei fusing prior to fertilization.Antipodal cells formed; 3; not proliferating; ephemeral. Synergids hooked(sometimes with filiform apparatus). Endosperm formation nuclear. Embryogenyonagrad.

Fruit non-fleshy; dehiscent (usually), or indehiscent (Cakile,Crambe,Coronopus etc), or a schizocarp (e.g. sometimes inCoronopus), or lomentaceous (or sometimes partially so). Mericarps whenschizocarpic, 2.Fruita silicula to a siliqua (with variations onthese themes, sometimes partially of entirely lomentaceous), orcapsular-indehiscent (i.e. when neither dehiscent nor lomentaceous).Capsules when dehiscent, valvular (the valves usually falling or spreadingupwards to reveal the replum). Seeds scantily endospermic, or non-endospermic;small to medium sized; winged (e.g.Matthiola,Isatis), orwingless. Embryo well differentiated. Cotyledons 2; flat, or folded(longitudinally or transversely), or rolled and twisted (e.g.inHeliophila). Embryo chlorophyllous (32/52); bent (nearly always folded,cotyledons against radicle), or straight (e.g.Leavenworthia), or otherthan straight, curved, bent or coiled (convolute and contorted inHeliophila). The radicle lateral, or dorsal.

Seedling.Germinationphanerocotylar, or cryptocotylar.

Physiology, phytochemistry.C₃ (almost exclusively), orC₃-C₄ intermediate. C₃ physiology recordeddirectly inBarbarea,Berteroa,Brassica,Cakile,Capsella,Cheiranthus,Crambe,Dipterygium,Farsetia,Matthiola,Raphanus,Sisymbrium,Thlaspi. C₃-C₄ intermediacy inMoricandia (3species).Anatomynon-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.).Mustard-oilspresent. Cyanogenic, or notcyanogenic. Alkaloids present, or absent (more often). Arbutin absent. Iridoidsnot detected. Proanthocyanidins absent (except sometimes in the seedcoat).Flavonols present, or absent; when present, kaempferol and quercetin. Ellagicacid absent (9 species, 7 genera). Aluminium accumulation not found. Sieve-tubeplastids P-type, or S-type; when P-type type I (b).

Geography, cytology.Frigid zoneto tropical. Cosmopolitan, concentrated in the North temperate andMediterranean.X = 5–12(+).

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 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,Marcus-kochia,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,Paysonia,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,Vesicaria,Warea,Weberbauera,Werdermannia,Winklera,Xerodraba,Yinshania,Zerdana,Zilla.

General remarks.For familyreview, see Vaughan, Macleod and Jones 1976.

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

Quotations.

When daisies pied,and violets blue,
And cuckoo-buds of yellow hue,
And Lady-smocks allsilver-white,
Do paint the meadows with delight
(‘Love’sLabour’s Lost’ —Cardamine pratensis)

Of paramours neraught he not a kers
(Chaucer, quoted by Ann Pratt 1857 via Jacobs,‘Flora of Cornwall’. Saxonkers = cress, employed to denotesomething worthless (i.e. referring to cruciferous weeds, rather thanwatercress); hence not caring a curse for something — with‘curse’ later becoming ‘damn’?)

In the season of thetulip cup,
When blossoms clothe the trees,
’Tis sweet to throw thelattice up
And scent thee on the breeze
(Moir, quoted by Ann Pratt,‘Wild Flowers’ (1857) - of Wallflowers)

And white and purplejiliflowers that stay
Lingering in blossom summer half away
And singleblood walls of a luscious smell
Old fashioned flowers which huswives love sowell
(John Clare 1827, ‘The Shepherd’s Calendar, June’ —Matthiola andCheiranthus

‘Shepheards Purse staiethbleeding in any part of the bodie, whether the iuice or the decoction thereof bedrunke, or whether it be used pultus wise or in bath or any other wise’.
(John Gerarde, ‘Herball’ 1597 —Capsellabursa-pastoris)

Illustrations.• Le Maout and Decaisne: Cheiranthus, Cochlearia,Parolinia. • Le Maout andDecaisne: Arabis, Barbarea, Lunaria, Tetrapoma (~Rorippa). • Le Maout and Decaisne: Cremolobus, Erophila,Eruca, Megacarpaea, Vesicaria (~Alyssoides). • Le Maout and Decaisne: Aethionema, Iberis, Isatis,Capsella, Lepidium, Neslia, Myagrum, Thlaspi. • Le Maout and Decaisne: Bunias, Clypeola, Crambe,Myagrum, Raphanistrum (~Raphanus), Thysanocarpus. • Aethionems grandiflorum (as A. cordifolium): Bot.Mag. 98 (1872). • Aethionemaspicatum: Hook. Ic. Pl. 15 (1885). • Alyssum samariferum: Hook. Ic. Pl. 15 (1885).• Anastatitica hierochuntica: Bot.Mag. 74 (1848). • Arabidopsisthaliana (as Draba thaliana): Eng. Bot. 115, 1863. • Arabis blepharophylla: Bot. Mag. 100 (1874).• Brassica napus: Köhler’s MedizinalPflanzen 1 (1887). • Brassicajuncea: Köhler’s Medizinal Pflanzen 1 (1887). • Brassica rapa (Turnip) and Brassica rapa ssp.campestris (Wild Turnip) : Eng. Bot. 89 and 90, 1863. • Brassica oleracea (Wild Cabbage) and Brassicanapus (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. • Cardamine kitaibelii (as Dentaria polyphylla):Bot. Mag. 111 (1885). • Cochlearia officinalis agg.: Köhler's MedizinalPflanzen 1 (1887). • Drabadedeana (as D. mawii): Bot. Mag. 101 (1875). • Erysimum perofskianum: Bot. Mag. 66 (1839).• Erophila verna (as 3 forms of Drabaverna): Eng. Bot. 134, 1863. • Farsetia burtonae: Hook. Ic. Pl. 14 (1880).• Heliophila scandens: Bot. Mag. 125(1899). • Isatis tinctoria: B.Ent. 734. • Leavenworthia aurea:Bot. Mag. 94 (1868). • Macropodium nivale: Bot. Mag. 67 (1840).• Marcus-kochia littorea (asMalcolmia): Bot. Mag. 78 (1852). • Matthiola fruticulosa (as M. coronopifolia): Bot.Mag. 126 (1900). • Matthiolasinuata var. oyensis: Bot. Mag. 126 (1900). • Moricandia arvensis: Bot. Mag. 57 (1830).• Moricandia moricandioides: Bot.Mag. 82 (1856). • Morisiamonanthes (as M. hypogaea): Bot. Mag. 124 (1898). • Orychophragmus violaceus (as Moricandiasonchifolia): Bot. Mag. 102 (1876). • Pachycladon novae-zelandii: Hook. Ic. Pl. 11(1867–71). • Paysoniagrandiflora (as Vesicaria): Bot. Mag. 63 (1836). • Physaria gracilis (as Vesicaria): Bot. Mag. 63(1836). • Rorippanasturtium-aquatica (Watercress, as Nasturtium officinalis): Eng. Bot. 125,1863. • Selenia aurea: Bot. Mag.108 (1882). • Sinapis alba (WhiteMustard): Köhler’s Medizinal-Pflanzen 3 (1898). • Streptanthus hyacinthoides: Bot. Mag. 63(1836). • Tchihatchewii isatidea:Bot. Mag. 124 (1898). • Thlaspirotundifolium (as Iberidella): Bot. Mag. 95 (1869). • Thysanocarpus curvipes, as T. elegans: Hook. Ic. Pl.1 (1837). • Brassiceae (Brassica,Diplotaxis: B. Ent. compilation, 1824–35). • Brassiceae (Rhynchosinapis, Cakile, Sinapis: B.Ent. compilation, 1824–35). • Lepidieae (Capsella, Coronopus, Cochlearia: B. Ent.compilation, 1824–35). • Lepidieae (Teesdalia), Drabeae (Erophila),Matthioleae (Matthiola): B. Ent. compilation, 1824–35. • Arabideae (Arabis, Cardamine, Barbarea: B. Ent.compilation, 1824–35). • Arabideae (Rorippa, Dentaria: B. Ent. compilation,1824–35). • Hesperideae(Erysimum, Cheiranthus, Hesperis: B. Ent. compilation, 1824–35).• Sisymbrieae (Arabidopsis,Descurainia, Alliaria: B. Ent. compilation, 1824–35). • Leaf hairs: Alyssum, Capsella, Cheiranthus,Hesperis, Matthiola (Solereder, 1908).

We advise against extracting comparative informationfrom the descriptions. This is much more easily achieved using theDELTA data files or theinteractive key, which allows access to the characterlist, illustrations, full and partial descriptions, diagnostic descriptions,differences and similarities between taxa, lists of taxa exhibiting or lackingspecified 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 alsoGuidelines for using data taken from Web publications.