The World Flora Online

SELECTED REFERENCES Al-Shehbaz, I. A. 1977. Protogyny in the Cruciferae. Syst. Bot. 2: 327-333. Al-Shehbaz, I. A. 1984. The tribes of Cruciferae (Brassicaceae) in the southeastern United States. J. Arnold Arbor. 65: 343-373. Al-Shehbaz, I. A. 1985. The genera of Brassiceae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 66: 279-351. Al-Shehbaz, I. A. 1985b. The genera of Thelypodieae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 66: 95-111. Al-Shehbaz, I. A. 1986. The genera of Lepidieae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 67: 265-311. Al-Shehbaz, I. A. 1987. The genera of Alysseae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 68: 185-240. Al-Shehbaz, I. A. 1988. The genera of Arabideae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 69: 85-166. Al-Shehbaz, I. A. 1988b. The genera of Anchonieae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 69: 193-212. Al-Shehbaz, I. A. 1988c. The genera of Sisymbrieae (Cruciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 69: 213-237. Al-Shehbaz, I. A., M. A. Beilstein, and E. A. Kellogg. 2006. Systematics and phylogeny of the Brassicaceae (Cruciferae): An overview. Pl. Syst. Evol. 259: 89-120. Al-Shehbaz, I. A., S. L. O’Kane, and R. A. Price. 1999. Generic placement of species excluded from Arabidopsis. Novon 9: 296-307. Al-Shehbaz, I. A. and S. I. Warwick. 2007. Two new tribes (Dontostemoneae and Malcolmieae) in the Brassicaceae (Cruciferae). Harvard Pap. Bot. 12: 429-433. Appel, O. and I. A. Al-Shehbaz. 2003. Cruciferae. In: K. Kubitzki et al., eds. 1990+. The Families and Genera of Vascular Plants. 9+ vols. Berlin etc. Vol. 5, pp. 75-174. Bailey, C. D. et al. 2006. Toward a global phylogeny of the Brassicaceae. Molec. Biol. Evol. 23: 2142-2160. Bailey, C. D., R. A. Price, and J. J. Doyle. 2002. Systematics of the halimolobine Brassicaceae: Evidence from three loci and morphology. Syst. Bot. 27: 318-332. Bailey, C. D., I. A. Al-Shehbaz, and G. Rajanikanth. 2007. Generic limits in the tribe Halimolobeae and the description of the new genus Exhalimolobos (Brassicaceae). Syst. Bot. 32: 140-156. Beilstein, M. A., I. A. Al-Shehbaz, and E. A. Kellogg. 2006. Brassicaceae phylogeny and trichome evolution. Amer. J. Bot. 93: 607-619. Beilstein, M. A., I. A. Al-Shehbaz, S. Mathews, and E. A. Kellogg. 2008. Brassicaceae phylogeny inferred from phytochrome A and ndhF sequence data: Tribes and trichomes revisited. Amer. J. Bot. 95: 1307-1327. Bowman, J. L. 2006. Molecules and morphology: Comparative developmental genetics of the Brassicaceae. Pl. Syst. Evol. 259: 199-215. German, D. A. and I. A. Al-Shehbaz. 2008. Five additional tribes (Aphragmeae, Biscutelleae, Calepineae, Conringieae, and Erysimeae) in the Brassicaceae (Cruciferae). Harvard Pap. Bot. 13: 165-170. Hall, J. C., K. J. Sytsma, and H. H. Iltis. 2002. Phylogeny of Capparaceae and Brassicaceae based on chloroplast sequence data. Amer. J. Bot. 89: 1826-1842. Hauser, L. A. and T. J. Crovello. 1982. Numerical analysis of generic relationships in Thelypodieae (Brassicaceae). Syst. Bot. 7: 249-268. Janchen, E. 1942. Das System der Cruciferen. Oesterr. Bot. Z. 91: 1-18. Koch, M. 2003. Molecular phylogenetics, evolution and population biology in Brassicaceae. In: A. K. Sharma and A. Sharma, eds. 2003+. Plant Genome: Biodiversity and Evolution. 2+ vols. in parts. Enfield, N. H. Vol. 1, part A, pp. 1-35. Koch, M. et al. 1999b. Molecular systematics of Arabidopsis and Arabis. Pl. Biol. (Stuttgart) 1: 529-537. Koch, M. et al. 2003b. Molecular systematics, evolution, and population biology in the mustard family (Brassicaceae). Ann. Missouri Bot. Gard. 90: 151-171. Koch, M., B. Haubold, and T. Mitchell-Olds. 2000. Comparative analysis of chalcone synthase and alcohol dehydrogenase loci in Arab

Herbs or subshrubs [shrubs or, rarely, lianas or trees], annual, biennial, or perennial; usually terrestrial, rarely submerged aquatics; with pungent watery juice; scapose or not; pubescent or glabrous, usually without papillae or tubercles (multicellular glandular papillae or tubercles present in Bunias, Chorispora, and Parrya); taprooted or rhizomatous (rarely stoloniferous), caudex simple or branched, sometimes woody, rhizomes slender or thick. Trichomes unicellular, simple, stalked, or sessile; forked, stellate, dendritic, malpighiaceous (medifixed, 2-fid, appressed), or peltate and scalelike, eglandular. Stems (absent in Idahoa, sometimes Leavenworthia) usually erect, sometimes ascending, descending, prostrate, decumbent, or procumbent; branched or unbranched. Leaves (sometimes persistent) cauline usually present, basal present or not (sometimes rhizomal present in Cardamine), rosulate or not, usually alternate (sometimes opposite or whorled in Cardamine angustata, C. concatenata, and C. diphylla and in Lunaria annua; sometimes subopposite in C. dissecta and C. maxima and in Draba ogilviensis), usually simple, rarely trifoliolate or pinnately, palmately, or bipinnately compound; stipules absent [with tiny, stipulelike glands at base of petioles and pedicels]; petiolate, sessile, or subsessile (sessile auriculate or not, sometimes amplexicaul); blade margins entire, dentate, crenate, sinuate, repand, or dissected. Inflorescences terminal, usually racemose (racemes often corymbose or paniculate) or flowers solitary on pedicels from axils of rosette leaves; bracts usually absent, sometimes present. Pedicels present (persistent or caducous [rarely geotropic]). Flowers bisexual [unisexual], usually actinomorphic (zygomorphic in Iberis, sometimes in Pennellia, Streptanthus, and Teesdalia); perianth and androecium hypogynous; sepals usually caducous, rarely persistent, 4, in 2 decussate pairs (1 pair lateral, 1 median), distinct [connate], not saccate or lateral (inner) pair (or, rarely, both pairs) saccate, forming tubular, campanulate, or urceolate calyx; petals 4, alternate with sepals, usually cruciform, rarely in abaxial and adaxial pairs, rarely rudimentary or absent, claw differentiated or not from blade, blade sometimes reduced and much smaller than well-developed claw, basally unappendaged, or, rarely, appendaged, margins entire or emarginate to 2-fid, rarely pinnatifid [fimbriate or filiform]; stamens (2 or 4) 6 [8-24], in 2 whorls, usually tetradynamous (lateral outer pair shorter than median inner 2 pairs), rarely equal in length or in 3 pairs of unequal length; filaments (slender, sometimes winged, appendaged, or toothed): median pairs usually distinct, rarely connate; anthers dithecal, dehiscing by longitudinal slits, pollen grains 3(-11)-colpate, trinucleate; nectar glands receptacular, variable in number, shape, size, and disposition around filament base, always present opposite bases of lateral filaments, median glands present or absent; disc absent; pistil 1, 2-carpellate; ovary 2-locular with false septum connecting 2 placentae, rarely 1-locular and eseptate, placentation usually parietal, rarely apical; gynophore usually absent; style 1, persistent [caducous], sometimes obsolete or absent; stigma capitate or conical, entire or 2-lobed, lobes spreading or connivent, sometimes decurrent, distinct or connate, rarely elongated into horns or spines; ovules 1-300 per ovary, anatropous or campylotropous, bitegmic, usually crassinucellate, rarely tenuinucellate. Fruits usually capsular, usually 2-valved ((3 or) 4(-6) in Rorippa barbareifolia, (2 or) 4 in Tropidocarpum capparideum), termed siliques if length 3+ times width, or silicles if length less than 3 times width, sometimes nutletlike, lomentaceous, samaroid, or schizocarpic and [with] without a c

Herbes , sous-arbrisseaux ou arbrisseaux. Poils presque toujours unicellulaires, simples ou ramifiés. Feuilles alternes sans stipules, ± nettement pétiolées, entières ou divisées, les basales souvent disposées en rosette. Inflorescences en grappes terminales ou parfois axillaires, avec ou sans bractées, souvent condensées et corymbiformes; infrutescences en grappes allongées. Fleurs bisexuées, régulières ou légèrement zygomorphes, tétramères, hypogynes, hétérochlamydées. Sépales 4, en 2 verticilles, libres, les 2 plus internes souvent sacciformes. Pétales 4 (rarement réduits), libres, ordinairement égaux, ± nettement onguiculés. Étamines 6 (rarement 4 ou 2), tétradynames, ordinairement libres; anthères biloculaires à déhiscence longitudinale; pollen trinucléé, le plus souvent 3-4- ou multicolpé. Nectaires entourant les filets à leur base, disposés sans ordre. Ovaire supère, sessile ou stipité, uniloculaire à placentation pariétale, formé de 2 carpelles soudés séparés par une fausse cloison membraneuse. Ovules généralement nombreux, anaou campylotropes. Stigmates bifides ou complètement soudés. Fruit : silique ou silicule déhiscente en deux valves, parfois akène ou fruit divisé en articles. Graines presqu'exalbuminées, cotylédons accombants, incombants ou diversement pliés.

Plantes herbacées'annuelles, bisannuelles ou vivaces.'Feuilles'alternes, rarement les inférieures opposées, exstipulées, simples, entières, dentées, profondément divisées ou composées, les supérieures souvent auriculées.'Fleurs'☿, actinomorphes, en racèmes généralement dépourvus de bractées ou solitaires et axillaires; sépales 4, en 2 verticilles; pétales 4, parfois 0; étamines 6, rarement 4 ou 2, hypogynes, tétradynames, en 2 verticilles, les 2 externes courtes et opposées aux sépales internes, les 4 internes longues et opposées par paires aux sépales externes, à filet mince ou ± élargi vers la base et à anthères basifixes et introrses; nectaires ± développés, réunis en disque ou isolés; gynécée à 2 carpelles, très rarement à 3-4; ovaire supère, sessile ou ± stipité, à placentas pariétaux, généralement 2-loculaire et muni d'un replum; style simple, à stigmate indivis ou 2-lobé.'Siliques'ou'silicules'déhiscentes en 2 valves s'écartant à maturité du replum, ou biarticulées : article supérieur nuciforme, caduc et indéhiscent; article inférieur stipitiforme et persistant.'Graines à albumen rare ou nul; embryon courbe; cotylédons plans ou condupliqués; radicule dorsale ou latérale.\n\t\t\tFamille cosmopolite, comprenant 350 genres groupant ± 3000 espèces, surtout abondamment représentée dans les régions extratropicales de l'hémisphère boréal et dont les quelques représentants tropicaux sont surtout des orophiles. Certaines espèces rudérales ont une aire de dispersion très étendue et croissent dans les stations les plus diverses.

Annual, biennial or perennial herbs (rarely somewhat shrubby) with alternate (rarely opposite or verticillate) exstipulate, simple or compound leaves sometimes forming a basal rosette Inflorescence usually racemose Fruit usually a dehiscent silique or silicule (more rarely indehiscent or transversely or longitudinally jointed) Seeds 1 to numerous, with no or very little endosperm Flowers actinomorphic (except for the stamens) usually bisexual, hypogynous Stamens usually 6, tetradynamous (rarely fewer or numerous); anthers 2- (rarely 1-) thecous, opening lengthwise Ovary sessile (rarely stipitate), syncarpous of 2 carpels, 1-locular with 1–2 parietal placentas or divided into 2 chambers by a false septum Sepals 4, free, in two series, often somewhat saccate Petals 4 (rarely fewer or absent) Inflorescence usually racemose Fruit usually a dehiscent silique or silicule (more rarely indehiscent or transversely or longitudinally jointed) Seeds 1 to numerous, with no or very little endosperm Flowers actinomorphic (except for the stamens) usually bisexual, hypogynous Stamens usually 6, tetradynamous (rarely fewer or numerous); anthers 2- (rarely 1-) thecous, opening lengthwise Ovary sessile (rarely stipitate), syncarpous of 2 carpels, 1-locular with 1–2 parietal placentas or divided into 2 chambers by a false septum Sepals 4, free, in two series, often somewhat saccate Petals 4 (rarely fewer or absent)

Fls typically borne in terminal, elongating, mostly bractless racemes (these occasionally with a few long branches), or rarely terminal and solitary on a scape, hypogynous, perfect, regular or nearly so, only seldom (as in Lunaria) with an evident gynophore; sep 4; pet 4, diagonal to the sep, commonly with an elongate claw and an abruptly spreading blade, collectively forming a cross, or seldom wanting; androecium tetradynamous, with 6 stamens, the 2 outer shorter than the 4 inner, or seldom the stamens fewer than 6; ovary bicarpellate, nearly always divided into 2 locules by a thin, unvasculated septum connecting the 2 parietal placentas, the placental frame forming a persistent replum; ovules (1–) several or many on each placenta, in 2 rows separated by the partition; style solitary, often short (or the stigma sessile); fr dry and usually dehiscent, generally a silique (elongate) or a silicle (short), the valves falling away from the persistent replum at maturity, or seldom the fr indehiscent and sometimes transversely jointed; endosperm very scanty or none; plants producing mustard-oils; herbs (all ours) or rarely shrubs, with alternate (rarely opposite), simple to often pinnately ± dissected lvs, only seldom with definite lfls. (Cruciferae) The following strictly artificial key avoids the more technical characters of the staminal glands, stigma, and embryo. 340/3350, mostly temp reg.

Herbs annual, biennial, or perennial, sometimes subshrubs or shrubs, with a pungent, watery juice. Eglandular trichomes unicellular, simple, stalked or sessile, 2- to many forked, stellate, dendritic, or malpighiaceous (medifixed, bifid, appressed), rarely peltate and scalelike; glandular trichomes multicellular, with uniseriate or multiseriate stalk. Stems erect, ascending, or prostrate, sometimes absent. Leaves exstipulate, simple, entire or variously pinnately dissected, rarely trifoliolate or pinnately, palmately, or bipinnately compound; basal leaf rosette present or absent; cauline leaves almost always alternate, rarely opposite or whorled, petiolate or sessile, sometimes absent. Inflorescence bracteate or ebracteate racemes, corymbs, or panicles, sometimes flowers solitary on long pedicels originating from axils of rosette leaves. Flowers hypogynous, mostly actinomorphic. Sepals 4, in 2 decussate pairs, free or rarely united, not saccate or lateral (inner) pair saccate. Petals 4, alternate with sepals, arranged in the form of a cross (cruciform; hence the earlier family name Cruciferae), rarely rudimentary or absent. Stamens 6, in 2 whorls, tetradynamous (lateral (outer) pair shorter than median (inner) 2 pairs), rarely equal or in 3 pairs of unequal length, sometimes stamens 2 or 4, very rarely 8-24; filaments slender, winged, or appendaged, median pairs free or rarely united; anthers dithecal, dehiscing by longitudinal slits. Pollen grains 3-colpate, trinucleate. Nectar glands receptacular, highly diversified in number, shape, size, and disposition around base of filaments, always present opposite bases of lateral filaments, median glands present or absent. Pistil 2-carpelled; ovary superior, sessile or borne on a distinct gynophore, mostly 2-locular and with a false septum connecting 2 placentae; placentation parietal, rarely apical; ovules anatropous or campylotropous, bitegmic, 1 to many per locule. Fruit typically a 2-valved capsule, generally termed silique (siliqua) when length 3 × or more than width, or silicle (silicula) when length less than 3 × width, dehiscent or indehiscent, sometimes schizocarpic, nutletlike, lomentaceous, or samaroid, segmented or not, terete, angled, or flattened parallel to septum (latiseptate) or at a right angle to septum (angustiseptate); valves 2(or 3-6); replum (persistent placenta) rounded, rarely flattened or winged; septum complete, perforated, reduced to a rim, or lacking; style 1, distinct, obsolete, or absent; stigma capitate or conical, entire or 2-lobed, sometimes lobes decurrent and free or connate. Seeds without endosperm, uniseriately or biseriately arranged in each locule, aseriate when 1, winged or wingless, mucilaginous or not when wetted; cotyledons incumbent (embryo notorrhizal: radicle lying along back of 1 cotyledon), accumbent (embryo pleurorrhizal: radicle applied to margins of both cotyledons), or conduplicate (embryo orthoplocal: cotyledons folded longitudinally around radicle), rarely spirally coiled (embryo spirolobal). Germination epigeal.

Ovary superior, sessile or stipitate, of seemingly 2 united carpels, divided into 2 “secondary” locules by a thin membranous septum (sometimes transversely locular); placentation parietal; ovules usually many, ana- or campylotropous; stigma bifid or connate Sepals 4, free, imbricate, the inner ones quite often saccate Petals 4 (rarely absent), free, usually equal, ± clawed, imbricate or contorted Stamens 6, tetradynamous (rarely 4 or 2), usually free; anthers usually 2-thecous, opening lengthwise; pollen grains mostly 3-, 4- or multicolpate Nectarial glands variously arranged at the filament-bases Herbs, sometimes subshrubs (very rarely small trees) Leaves alternate, basal ones often in a rosette, not stipulate, ± petiolate, entire to variously divided Inflorescences terminal or sometimes axillary racemes, in flower mostly condensed and often corymbose, in fruit elongate, usually ebracteate Flowers hermaphrodite, actinomorphic or slightly zygomorphic, hypogynous, 4-merous, heterochlamydous Fruit a bivalved dehiscent siliqua or silicula (see key), sometimes a nutlet, lomentum or otherwise constructed Seeds virtually devoid of endosperm, with cotyledons incumbent, accumbent or variously folded Sepals 4, free, imbricate, the inner ones quite often saccate Petals 4 (rarely absent), free, usually equal, ± clawed, imbricate or contorted Stamens 6, tetradynamous (rarely 4 or 2), usually free; anthers usually 2-thecous, opening lengthwise; pollen grains mostly 3-, 4- or multicolpate Nectarial glands variously arranged at the filament-bases Herbs, sometimes subshrubs (very rarely small trees) Leaves alternate, basal ones often in a rosette, not stipulate, ± petiolate, entire to variously divided Inflorescences terminal or sometimes axillary racemes, in flower mostly condensed and often corymbose, in fruit elongate, usually ebracteate Flowers hermaphrodite, actinomorphic or slightly zygomorphic, hypogynous, 4-merous, heterochlamydous Fruit a bivalved dehiscent siliqua or silicula (see key), sometimes a nutlet, lomentum or otherwise constructed Seeds virtually devoid of endosperm, with cotyledons incumbent, accumbent or variously folded