CARDAMINE Linnaeus, Sp. Pl. 2: 654. 1753. Ihsan A
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CARDAMINE Linnaeus, Sp. Pl. 2: 654. 1753. Ihsan A. Al-Shehbaz and Karol Marhold Tribe: Cardamineae Dumort., Fl. Belg.: 124. 1827. Name derivation: Greek kardamon, used by Dioscorides for some species of Brassicaceae. Lectotype species (designated by Britton & Brown, Ill. Fl. N.U.S., ed. 2, 2: 183. 1913): C. pratensis L. Dentaria Linnaeus, Sp. Pl. 2: 653. 1753. Lectotype species (designated by Britton & Brown, Ill. Fl. N.U.S., ed. 2, 2: 187. 1913: D. pentaphyllos L. Dracamine Nieuwland, Amer. Midl. Naturalist 4: 40. 1915. Type species: not designated. Heterocarpus Philippi, Bot. Zeitung 14: 641. 1856, not Wight (1853). Type species: H. fernandezianus R. A. Philippi. Ghinia Bubani, Fl. Pyrenaea 3: 158. 1901, non Schreber (1789). Type species: not designated. Iti Garnock-Jones & P. N. Johnson, New Zealand J. Bot. 25: 603. 1988. Type species: I. lacustris Garnock-Jones & P. N. Johnson. Loxostemon J. D. Hooker & Thomson, J. Proc. Linn. Soc., Bot. 5: 129. 1861. Type species: L. pulchellus J. D. Hooker & Thomson. Porphyrocodon W. J. Hooker in Bentham & J. D. Hooker, Gen. Pl. 1: 79. 1862. Type species: P. pictus (W. J. Hooker) B. D. Jackson. Pteroneurum de Candolle, Mém. Mus. Hist. Nat. 7: 231. 1821. Type species: not designated. Sphaerotorrhiza (O. E. Schulz) Khokhrjakov, Fl. Magadansk. Obl. 235. 1985. Type species: S. trifida (Poiret ex Lamarck) Khokhrjakov. Herbs, annual, biennial, or perennial, rhizomatous or tuberous, sometimes with axillary bulbils. Trichomes absent or simple. Multicellular glands absent. Stems erect or prostrate, leafy or rarely leafless and plant scapose. Basal leaves petiolate, rosulate or not, simple and entire, toothed, or 1–3-pinnatisect, or palmately lobed, sometimes trifoliolate, pinnately, palmately, or bipinnately compound; cauline leaves alternate, rarely opposite or whorld, simple or compound as basal leaves, petiolate or sessile and base cuneate to attenuate or auriculate to sagittate, margin entire, dentate, or variously lobed. Racemes several to many-flowered, ebracteate or rarely bracteate throughout, corymbose or in panicles, elongated in fruit; rachis straight or rarely flexuous; fruiting pedicels slender or thickened, erect, divaricate, or reflexed. Sepals ovate or oblong, free, caducous or rarely persistent, erect to rarely spreading, equal or unequal, base of lateral pair saccate or not. Petals white, pink, purple, or violet, never yellow, erect to spreading, longer than sepals, rarely absent; blade obovate, spatulate, oblong, or oblanceolate, apex obtuse or emarginate; claw absent or strongly differentiated from blade, longer to shorter than sepals, glabrous, unappendaged, entire. Stamens 6 and slightly to strongly tetradynamous, rarely 4 and equal in length, erect or spreading, exserted or included; filaments wingless, unappendaged, glabrous, free, dilated or not at base; anthers ovate to oblong or linear, not apiculate at apex. Nectar glands confluent and subtending bases of all stamens; median nectaries 2 or rarely 4 or absent; lateral nectaries annular or semiannular. Ovules 4−80 per ovary; placentation parietal. Fruit dehiscent capsular siliques, linear or rarely oblong, elliptic, or narrowly lanceolate, slightly to strongly latiseptate, not inflated, unsegmented; valves papery, not veined or rarely proximal part with an obscure midvein, glabrous or very rarely hairy, not keeled, flat, not covering entire fruit width, smooth or torulose, wingless, unappendaged, dehiscing elastically acropetally, spirally or circinately coiled; gynophore absent; replum strongly flattened, visible; septum complete, membranous, tansluscent, veinless; style distinct and slender, rarely obsolete, persistent, glabrous; stigma capitate, entire, unappendaged. Seeds 1 uniseriate, wingless or not margined, oblong to ovate, slightly flattened; seed coat smooth, minutely reticulate, colliculate, or rugose; mucilaginous or not when wetted; cotyledons accumbent or very rarely incumbent. × = 7, 8. 256 species: worldwide and with native species on all continents except Antarctica. References: Interactive key to the genera is being prepared 2 acris diphylla lineariloba purpurascens africana dissecta lojanensis purpurea altigena douglasii longii quinquefolia anemonoides elegantula longipedicellata raphanifolia angulata engleriana loxostemonoides repens angustata enneaphyllos luxurians resedifolia anhuiensis enriquei lyrata rivularis amara eremita macrocarpa robusta apennina fallax macrophylla rockii appendiculata fargesiana majovskii rostrata arakiana flagellifera marginata rotundifolia armoracioides flexuosa maritima rupicola asarifolia fragariifolia matthioli scaposa astoniae franchetiana maxima schinziana auriculata franklinensis mexicana schulzii balnearia fulcrata micranthera scutata bellidifolia gallaecica microphylla seidlitziana bilobata georgiana microthrix seravschanica blaisdellii geraniifolia microzyga silana bodinieri glacialis moirensis simplex bonariensis glanduligera monteluccii speciosa breweri glauca multiflora sphenophylla bulbifera glechomifolia multijuga stellata bulbosa gouldii nepalensis stenoloba calcicola gracilis niigatensis subcarnosa caldeirarum graeca nipponica tanakae californica granulifera nuttallii tangutorum calthifolia granulosa nymanii tenera carnosa griffithii obliqua tenuifolia caroides gunnii occidentalis tenuirostris castellana heptaphylla oligosperma tianqingiae changbaiana hirsuta opizii trichocarpa chelidonia hispidula ovata trifida chenopodiifolia holmgrenii pachystigma trifolia cheotaiyienii hydrocotyloides papillata trifoliolata chilensis hygrophila papuana tryssa circaeoides iliciana parviflora tuberosa clematitis impatiens pattersonii uliginosa concatenata jamesonii paucifolia uniflora conferta jejuna paucijuga umbellata constancei jonselliana pectinata variabilis cordata keysseri pedata victoris cordifolia kitaibelii penduliflora violacea corymbosa komarovii pensylvanica volckmannii crassifolia kruesselii pentaphyllos vulgaris debilis lacustris penzesii waldsteinii delavayi latior picta yezoensis densiflora lazica plumieri yunnanensis dentata leucantha pratensis depressa lihengiana prorepens digitata lilacina pulchella Species in red are not yet described 3 Cardamine acris Grisebach, Spicil. Fl. Rumel. 1: 253. 1843; C. raphanifolia Pourret subsp. acris (Grisebach) O. E. Schulz, Bot. Jahrb. Syst. 32: 512. 1903. TYPE: [F.Y.R. Makedonija] “gregarie ad fontem m. Kobelitza [Kobelica] juxta Mandram Weitzensem alt. 4200′, 21 Jul 1839, Grisebach 943 (lectotype designated by Strid (1986: 257), GOET; isolectotype, K). Cardamine africana Linnaeus, Sp. Pl. 2: 655. 1973. Lectotype (designated by Marais, 1970): Hermann, Paradisus Batavus Fig. p. 202 (1698). Cardamine innovans O. E. Schulz, Bot. Jahrb. Syst. 32: 417. 1903. TYPE: Guatemala, above Tecpam, 2500 m, 1882, F. C. Lehmann 1475 (lectotype, here designated, G-BOIS). Cardamine borbonica Persoon, Syn. Pl. 2: 195. 1807. TYPE: Réunion [Bourbon], Bory de St. Vincent (holotype, G). Cardamine holtziana Engler & O. E. Schulz, Bot. Jahrb. Syst. 32: 416. 1903. TYPE: Tanzania, Lushoto District, W. Usambara Mts, between Wambugu and Malalo, A. Engler 1356 (holotype, B). Cardamine ovata Bentham var. corymbosa Britton, Bull. Torrey Bot. Club 16: 16. 1889. TYPE. Bolivia, [La Paz], Undavi, 10000 ft, Oct 1885, H. H. Rusby 1206 (holotype, NY!). Herbs, perennial. Rhizomes slender, not tuberous. Stems (5−)25−70(−80) cm, erect to ascending, simple or sometimes branched near middle, glabrous throughout or pubescent. Basal leaves not rosulate; lower and middle cauline leaves 4−10(−15) cm, trifoliolate; petiole 2−6(−7) cm; terminal leaflet (1−)1.5−7 × 1.5−4 cm, lanceolate to ovate, sparsely strigose with trichomes to 0.6 mm, base cuneate to subobtuse, margin ciliolate or glabrous, serrate or crenate, teeth mucronate, apex acute, petiolule 2−2.5 cm; lateral leaflets same size or slightly smaller than terminal one, base oblique. Racemes ebracteate, slightly elongated or not elongated in fruit; rachis straight; fruiting pedicels 0.5−3 cm, ascending, stout, straight. Sepals oblong, 1.5−4 × 1−1.5 mm, not saccate, caducous, glabrous; petals white or lavender, spatulate, 3.5−9 × 2−4 mm, apex obtuse; filaments 2.5−5 mm; anthers oblong, 1−1.5 mm; ovules 8−20 per ovary. Fruits linear, (2.5−)3.5−5(−6) cm × 1.5−2.5 mm, glabrous; style 0.5−1.5 mm, stout. Seeds brown, oblong, 1.5−2.5 × 1−1.5 mm. Flowering: Sep−Jan. Habitat: Moist or wet areas, mixed forests, disturbed places, roadside clearings, damp shady areas. Elevation: 500−3400 m. Distribution: native of Africa (Comores, Ethiopia, Kenya, Madagascar, South Africa, Tanzania, Ugunda), naturalized in Asia (India, Indonesia, Malesia, Papua New Guinea, Sri Lanka), Central America (Costa Rica, El Salvador, Guatemala, Honduras, Mexico [Chiapas]), and South America [Argentina (Misiones, Salta, Tucumán), Bolivia (Cochabamba, La Paz, Santa Cruz, Tarija), Brazil (Parana, Río Grande Do Sul, Santa Catalina), Columbia (Cundinamarca), Ecuador (Santiago-Zamora), Peru (Amazonas, Cajamarca, Cuzco), Venezuela (Mérida)]. Specimens examined: ARGENTINA. Misiones: Gral. Manuael Belgrano, 8 km de Bernardo de Irigoyen hacia San Antonio, 26°12′S, 53°40′W, Morrone et al. 1378 (MO, SI). Salta: Santa Victoria, camino al Angosto del Baritú, 22°30′S, 64°45′W, Ahumada & Agüero 8343 (SI); Parq Nac. Baritú, Lipeo, Rio Naranjo, Zuloaga et al. 1165 (BACP, SI); Anta, Parque Nac. El Rey, Picada al Chomo Los Loios, Ezcurra, Ponce & Brown 431 (SI). Dep. Anta, Res. Nac. Fca. Del Rey, Ezcurra et al. 431 (BACP). Tucumán: Dep. Chicligasta, arroyo