BRASSICALES Bromhead Main Tree, Synapomorphies

Brassicales Bromhead in Edinburgh New Philos. J. 24: 416. Apr 1838; Capparanae Reveal in Phytologia 76: 3. 2 Mai 1994

Fossils Dressiantha bicarpellata from the Turonian Raritan Formation in New Jersey (late Cretaceous) comprises somewhat zygomorphic flowers with four sepals, five petals, five antepetalous stamens with disporangiate monothecal anthers containing tricolporate pollen grains, a ring of basally connate alternipetalous staminodia, and two basally connate carpels inserted on top of a well developed gynophore.

Habit Usually bisexual (sometimes monoecious, andromonoecious, gynomonoecious, polygamomonoecious, dioecious, or polygamodioecious), evergreen or deciduous trees, shrubs or suffrutices, or perennial, biennial or annual herbs (rarely lianas).

Vegetative anatomy Phellogen ab initio superficially or deeply seated (subepidermal or pericyclic), or absent. Secondary lateral growth usually normal (sometimes anomalous, from concentric cambia) or absent. Vessel elements with simple or scalariform (rarely reticulate) perforation plates; lateral pits usually alternate (rarely opposite or scattered), simple or bordered pits. Vestured pits often present. Imperforate tracheary xylem elements usually fibre tracheids or libriform fibres (sometimes tracheids) with simple or bordered pits, septate or non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse (sometimes diffuse-in-aggregates), or paratracheal scanty vasicentric, confluent, or banded (rarely aliform, lozenge-aliform or confluent). Wood elements often storied. Intraxylary phloem rarely present. Sieve tube plastids usually S₀ or Ss type (rarely Pcs type). Nodes 1:1, 1:2, 2:2 or ≥3:≥3, unilacunar to trilacunar (rarely multilacunar), with one or three leaf traces (rarely two or more traces). Laticifers sometimes present. Calciumcarbonate, calciumsulfate and/or calciumoxalate crystals (as prismatic crystals, sometimes druses) often frequent (sometimes also silica).

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple, furcate or many-branched (occasionally T-shaped malpighiaceous hairs), stalked or unstalked, medifixed (with two branches parallel to epidermis), stellate (with several or many branches, from common point, parallel to epidermis), candelabra-shaped, dendritic, peltate or lepidote (rarely prickles), or absent; glands or glandular hairs unicellular or multicellular, uniseriate, simple (occasionally peltate, sometimes with calciumcarbonate), or absent.

Leaves Alternate (usually spiral, rarely distichous; rarely opposite), simple or pinnately or palmately compound (sometimes twice or three times compound), entire or pinnately or palmately lobed, sometimes coriaceous or succulent (rarely almost absent), with conduplicate, supervolute, involute, flat or curved ptyxis. Stipules intrapetiolar or cauline (sometimes replaced by glands, extrafloral nectaries or spines), or absent; leaf sheath absent. Petiole vascular bundle transection arcuate or annular. Venation usually pinnate (sometimes palmate, rarely flabellate). Stomata usually anomocytic or anisocytic (sometimes paracytic, rarely helicocytic, cyclocytic, tetracytic, staurocytic, etc.). Cuticular wax crystalloids sometimes as rosettes of platelets (Fabales type) or tubuli, or absent. Stomatal (including guard cells) or idioblastic myrosin cells (with myrosinase) usually frequent. Mesophyll often with crystalliferous cells containing calciumoxalate crystals or druses, or with sclerenchymatous idioblasts containing dendrosclereids or other types of sclereids. Epidermis usually with mucilage cells; epidermal cells sometimes with crystals. Leaf margin serrate, sinuate or entire.

Inflorescence Terminal or axillary, corymb, panicle, thyrsoid, raceme, spike or umbel (rarely catkin), or solitary axillary. Bracts and floral prophylls (bracteoles) usually absent.

Flowers Actinomorphic, zygomorphic or bisymmetric. Hypanthium sometimes present. Usually hypogyny (rarely half epigyny). Receptacle in zygomorphic flowers sometimes with nectariferous appendage or gland inside adaxial sepal; often elongated into androgynophore or gynophore. Sepals (two to) four or five (to ten), usually with imbricate (sometimes valvate, contorted or open) aestivation, in one or two whorls, usually caducous, usually free (rarely connate or absent). Petals (two to) four to six (to nine), with imbricate, valvate or contorted (rarely open) aestivation, alternisepalous, often clawed, usually free (rarely absent). Nectariferous disc usually extrastaminal or intrastaminal, annular, or absent; nectariferous glands of various shape, inserted on disc, perianth or stamen, or absent. Disc present or absent.

Androecium Stamens usually four, five or 2+4 (sometimes eight to ten, rarely one, two or up to c. 250), in one or several whorls, usually haplostemonous or diplostemonous (rarely triplostemonous). Filaments usually in one whorl, usually free (sometimes more or less connate), usually free from tepals (sometimes adnate at base to petals, epipetalous). Anthers basifixed or dorsifixed, versatile or non-versatile, usually tetrasporangiate (rarely disporangiate), usually introrse (sometimes latrorse or extrorse), longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia usually absent (sometimes four to ten).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually 3–4-colpate (sometimes 3–4-colporate or 3–4-colporoidate, rarely 2–11-colpate, 2–11-colporate, hexacolporoidate or inaperturate), shed as monads, bicellular or tricellular at dispersal. Exine tectate or semitectate, usually with columellate (rarely acolumellate, sometimes granular) infratectum, perforate, finely punctate, reticulate, microreticulate, striate, foveolate, scabrate, spinulate or smooth.

Gynoecium Pistil composed of two to six (to eight) connate and often paracarpous carpels. Ventral carpellary vascular bundles fused and highly developed. Ovary usually superior (rarely semi-inferior), trilocular or ab initio unilocular, and later usually bilocular with membranous secondary septum (rarely quinquelocular to octalocular), usually sessile (sometimes stipitate, with gynophore). Style single, simple, bilobate or trilobate, or absent, or stylodia (four or) five free. Stigma one, capitate or lobate, or stigmas (four or) five, truncate or punctate (sometimes flabellate or almost petaloid), usually papillate, Dry type. Pistillodium usually absent (male flowers sometimes with pistillodium). Strongly developed fused ventral carpellary vascular bundles often present.

Ovules Placentation axile, apical or parietal (sometimes intrusively parietal, rarely basal-parietal, basal or laminar). Ovules one to more than 300 per carpel, in one or two rows, usually anatropous or campylotropous (rarely amphitropous), ascending, horizontal or pendulous, apotropous or epitropous, usually bitegmic (sometimes unitegmic), usually crassinucellar (rarely weakly crassinucellar to incompletely tenuinucellar). Micropyle usually bistomal (rarely endostomal), sometimes Z-shaped (zig-zag). Nucellar cap sometimes present. Hypostase often present. Archespore usually unicellar (sometimes bicellular or multicellular). Megagametophyte usually monosporous, Polygonum type (rarely disporous, Allium type, or modified Drusa type). Synergids sometimes with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustorium chalazal, lateral or absent. Embryogenesis usually onagrad (sometimes asterad, rarely solanad).

Fruit Usually a loculicidal and/or septicidal capsule (often dehiscing from base upwards by usually two valves, with membranous secondary septum and persistent replum consisting of placental tissue (sometimes a nut; rarely berry, pumpkin fruit, drupe, samara, schizocarp, drupaceous syncarp, or an assemblage of follicles).

Seeds Aril usually absent (elaiosome rarely present). Seed coat usually endotestal (sometimes exotegmic, rarely also mesotegmic). Testa sometimes vascularized and multiplicative (outer testal epidermis sometimes modified into mucilaginous sarcotesta). Exotestal cells sometimes palisade, thick-walled. Mesotestal cell walls sometimes sclerified or tanniniferous. Endotestal cell walls sometimes sclerified and crystalliferous. Tegmen often multiplicative. Exotegmen and mesotegmen usually crushed. Exotegmen sometimes (rarely also mesotegmen) sclerified, fibrous or non-fibrous. Endotegmic cell walls sometimes tanniniferous and lignified. Perisperm rarely developed. Endosperm copious or sparse, oily, or absent. Embryo curved, plicate or straight, well differentiated, oily, with or without chlorophyll. Cotyledons two (or three). Germination usually phanerocotylar (sometimes cryptocotylar).

Cytology x = (4–)8(–15) Endoplasmic reticulum with protein-rich dilated organelle-like cisternae.

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, delphinidin, ellagic and gallic acids, caffeic acid, oleanolic acid derivatives, proanthocyanidins (prodelphinidins), protoalkaloids, pyrrolidine alkaloids, piperidine alkaloids, glucosinolates (mustard oil glycosides with R-N=C=S group, i.e. 2-hydroxy-2-methylpropylic benzylglucosinolate, 3,4-dihydroxybenzyl glucosinolate, phenethylglucosinolate, etc.), derived from phenylalanine, tyrosine, methionine, valine, isoleucine and/or leucine, saponins, cyanogenic compounds, hydroxyproline betaines, mustard oils based on glucotropaeolin, glucocapparin, glucocleomin, cholinesters (sinapin, cochlearin, hesperalin, isoferuloylcholin, p-cumaroylcholin, etc.) etc., erucic acid, n-eicose-11-enoic acid, docosadienoic acid and other unsaturated fatty acids, sinapic acid, ferulic acid, benzylisothiocyanate, methoxybenzylisothiocyanates, cucurbitacins, myo-inositol, sweet-tasting proteins (brazzein, pentadin), and aromatic m-carboxycinnamic acids and proteolytic enzymes (papain, carpain) present. Myricetin, tannins and other polyphenolic compounds rare.

More than 130 different kinds of glucosinolates are known. Digested into glucose and aglucones by thioglucoside glucohydrolase (myrosinase, a β-thioglucohydrolase), when tissues are wounded. The aglucones are transformed into thiocyanates (mustard oils with R-N=C=S), toxic isothiocyanates and often toxic or non-toxic nitriles and other compounds.

Akaniaceae and Tropaeolaceae have the following potential synapomorphies in common (Stevens 2001 onwards): young stem with separate vascular bundles; vessel elements with scalariform perforation plates; axial parenchyma sparse vasicentric; absence of floral prophylls (bracteoles); flowers obliquely zygomorphic, large; presence of perianth tube or hypanthium; petals clawed; stamens eight, with short connective prolongations; style elongated; placentation apical or apical-axile; ovules one or two per carpel, epitropous; and testa vascularized.

Moringaceae and Caricaceae share the following potential synapomorphies (Stevens 2001 onwards): woody habit, with stout stems (often pachycaul); cambium storied; nodes also multilacunar; presence of ER-dependent vacuoles; stipules modified into glands; colleters present on petiole and/or lamina; leaf venation palmate; cuticular wax crystalloids as rosettes of platelets; inflorescence thyrse; flowers pentamerous, whitish; carpels antesepalous; ovary longitudinally sulcate; style hollow; placentation parietal, with placental vascular strands opposite ventral bundles; ovules numerous per carpel; micropyle bistomal; outer integument five or six cell layers thick; testa multiplicative; and mesotesta lignified.

The [Koeberliniaceae+[Bataceae+Salvadoraceae]] clade is characterized by the following features: absence of idioblastic myrosin cells; flowers tetramerous; pollen grains tricolporoidate; pistil composed of two connate carpels; fruit indehiscent; exotestal cells well developed; and x = 11. Adaptations to dry and/or salt environments are very common.

Bataceae and Salvadoraceae share the characteristics: storied wood; non-bordered perforation plates; wide, multiseriate wood rays; nodes 1:2, unilacunar with two traces; opposite leaves; bracts with apical colleters; secondary veins palmate, ascending from at or near leaf base; paracytic stomata; basal placentation; two ovules per carpel; non-fibrous exotegmen; absence of endosperm; and straight to somewhat curved embryo.

The clade [Emblingiaceae+[Pentadiplandraceae+[[Gyrostemonaceae+[Borthwickiaceae+Resedaceae]]+ Tovariaceae+Brassicaceae]] have the following potential synapomorphies, according to Stevens (2001 onwards): cisternae of endoplasmic reticulum dilated and vacuole-like; cuticular wax crystalloids absent; inflorescence terminal, racemose; floral prophylls (bracteoles) absent; floral development open; petals clawed; nectary extrastaminal; ovules inserted in two rows; endotesta crystalliferous; extension of 3’ terminus of plastid gene rbcL; glucosinolates synthesized also from valine/isoleucine and/or leucine and indole glucosinolates from tryptophane.

Gyrostemonaceae and Resedaceae share the synapomorphies: absence of idioblastic myrosin cells; unicellular hairs; presence of stylodia; calyx persistent in fruit; and presence of funicular aril.

Tiganophytaceae (Swanepoel & al. 2020). Tiganophyton (1; T. karasense; the arid Karas Region in southeastern Namibia).

Bretschneideraceae Engl. et Gilg, Engler’s Syllabus, ed. 9-10: 218. 6 Nov 1924, nom. cons.; Akaniales Doweld in Byull. Mosk. Obshch. Ispyt. Prir., Biol. 105(5): 60. 9 Oct 2000

Distribution Southeastern China, Vietnam, Taiwan, Thailand, eastern Australia.

Fossils Uncertain. Fossil leaves attributed to Akania have been found in Paleocene layers in Argentina.

Vegetative anatomy Phellogen ab initio subepidermal. Primary medullary strands usually wide. Young stem with separate vascular bundles. Vessel elements with simple and/or scalariform (sometimes reticulate) perforation plates; lateral pits alternate, simple and/or bordered pits. Imperforate tracheary xylem elements libriform fibres with simple (Bretschneidera) or bordered (Akania) pits, septate or non-septate. Wood rays multiseriate, heterocellular. Axial parenchyma paratracheal scanty vasicentric or banded. Wood elements not storied. Sieve tube plastids ? type (Akania) or P type (?) (Bretschneidera). Nodes 3:3, trilacunar with three leaf traces. Bark and inflorescences with idioblastic and stomatal myrosin cells (in Akania sometimes absent). Calciumoxalate in Akania as druses and single crystals.

Leaves Alternate (spiral), imparipinnate, in Akania large, coriaceous, with supervolute to curved ptyxis. Stipules small or absent; leaf sheath absent. Petiole vascular bundle transection? Petiolules swollen or articulated. Venation pinnate. Stomata in Akania present in groups, in Bretschneidera evenly dispersed. Cuticular waxes absent; cuticular cracks distinct. Leaflet margins spinose-serrate (Akania) or entire (Bretschneidera).

Inflorescence Terminal, raceme (Bretschneidera) or axillary panicle (Akania). Floral prophylls (bracteoles) absent.

Flowers Actinomorphic (Akania) or obliquely zygomorphic (Bretschneidera). Hypanthium or perianth tube cupular. Hypogyny or perigyny. Sepals five, with imbricate aestivation, free (Akania) or connate (Bretschneidera). Petals five, with contorted or imbricate aestivation, often clawed, inner petals smaller than outer, free. Nectariferous disc in Bretschneidera annular, intrastaminal; nectary and disc absent in Akania.

Androecium Stamens five outer and three to five inner or three (rarely four), diplostemonous; abaxial stamen in antepetalous whorl (Akania), in Bretschneidera bent down and inserted below one petal. Filaments hairy at base, inserted at receptacular base (in Bretschneidera at nectariferous disc), free from each other and from tepals. Anthers subbasifixed (Akania) or dorsifixed (Bretschneidera), versatile, tetrasporangiate, latrorse (Akania) or introrse (Bretschneidera), longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (rarely dicolpate), shed as monads, bicellular at dispersal. Exinc semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of three (or five) connate carpels. Ovary superior or semi-inferior, trilocular (or quinquelocular), stipitate (with gynophore). Style single, simple, narrow (in Bretschneidera bent down). Stigma small, capitate (Bretschneidera), trilobate (in Bretschneidera hollow), type? Pistillodium absent.

Ovules Placentation apical-axile. Ovules two (or three) per carpel, anatropous (Akania) or campylotropous (Bretschneidera), pendulous, epitropous?, bitegmic, crassinucellar. Micropyle bistomal. Outer integument approx. five cell layers thick, multiplicative, vascularized. Inner integument two or three cell layers thick. Hypostase present. Megagametophyte in Akania monosporous, Polygonum type; in Bretschneidera disporous, 8-nucleate, Allium type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Seeds Aril absent. Testa vascularized, multiplicative. Exotestal cells palisade, thick-walled. Mesotesta thick, with thick sclerified cell walls. Endotesta thickened. Tegmen not multiplicative, thin, more or less crushed. Exotegmen non-fibrous. Perisperm not developed. Endosperm copious(Akania) or absent (Bretschneidera). Embryo large, straight, well differentiated, chlorophyll? Cotyledons two, large, thick. Germination phanerocotylar.

Phytochemistry Flavonols (kaempferol, quercetin), cyanidin, proanthocyanidins (prodelphinidins), alkaloids, and glucosinolates (2-hydroxy-2-methylpropylic and 3,4-dihydroxybenzylic glucosinolate, synthesized from tyrosine, in Bretschneidera also from valine/isoleucine and/or leucine) present. Cyanogenic compounds? Ellagic acid and saponins not found.

Distribution Coasts of southern New Guinea and northern Australia, tropical and subtropical coasts in America and the Galápagos Islands.

Habit Dioecious (B. maritima) or monoecious (B. argillicola), evergreen suffrutices or shrubs. Succulent halophytes or xerophytes. Young stems quadrangular in cross-section.

Vegetative anatomy Phellogen ab initio in pericyclic fibre bundles. Secondary lateral growth often absent. Vessel elements with simple perforation plates (with rudimentary borders); lateral pits usually alternate (rarely opposite), bordered pits. Vestured pits present. Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate? Wood rays usually multiseriate, heterocellular. Axial parenchyma apotracheal or paratracheal scanty vasicentric. Wood elements (including parenchyma) sometimes storied. Sieve tube plastids S type. Nodes 2?:2, bilacunar? with two leaf traces. Myrosin cells stomatal; idioblastic myrosin cells absent. Crystals?

Leaves Opposite, simple, entire, fleshy, reduced, with ? ptyxis. Stipules very small, without vascular bundles, intrapetiolar or cauline, often caducous; leaf sheath absent. Leaf traces? disappearing adjacent to nodes; leaf supported by two vascular bundles from stem margins. Venation consisting of two bundles. Stomata paracytic or anomocytic. Cuticular wax crystalloids? Myrosin cells stomatal. Leaf margin entire.

Inflorescence Axillary, racemose, in Batis maritima cone-like dense catkin (strobilus); in Batis argillicola loose spike. Bracts in male inflorescences with cochleariform and imbricate aestivation; bracts in female inflorescence small and partially incorporated in fleshy spike. Floral prophylls (bracteoles) absent?

Flowers Zygomorphic (male flowers) or actinomorphic? (female flowers), minute. Female flowers in Batis maritima connate. Hypogyny? Sepals? (bracteoles?) in male flowers two, unequal in size (adaxial flowers larger than and overlapping abaxial flowers), median, enclosing flower (Batis argillicola), or four, connate (Batis maritima), absent in female flowers. Petals? in male flowers four, clawed, free or connate at base, absent in female flowers. Nectary absent. Disc absent.

Androecium Stamens four, haplostemonous, antesepalous?, alternipetalous. Filaments free from each other and from tepals. Anthers dorsifixed, versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); connective slightly prolonged apically. Tapetum secretory, with binucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains 3(–4)-colpor(oid)ate, shed as monads, bicellular at dispersal. Exine pertectate, with acolumellate infratectum and undifferentiated ectexine, microverrucate, almost smooth. Ectexine spongy, undifferentiated.

Gynoecium Pistil composed of two connate carpels; carpel divided by secondary septum. Ovary largely quadrilocular (unilocular at base); ovaries of adjacent flowers more or less connate and adnate to bract bases. Style very short or absent. Stigma capitate-penicillate to slightly bilobate, persistent, papillate, type? Male flowers often with rudimentary pistillodium.

Ovules Placentation basal-parietal. Ovules two per carpel, collateral, anatropous, ascending, epitropous, bitegmic, crassinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument ? cell layers thick. Inner integument ? cell layers thick. Nucellar cap present. Megagametophyte monosporous, Polygonum type? Endosperm development nuclear? Endosperm haustoria? Embryogenesis?

Fruit A drupaceous syncarp composed of fused flowers (Batis maritima), or a drupe with four one-seeded pyrenes (Batis argillicola). Endocarp lignified.

Seeds Aril absent. Seed coat exotestal? Testa membranous, multiplicative? Tegmen multiplicative? Exotegmen non-fibrous? Perisperm not developed. Endosperm absent. Embryo almost straight, well differentiated, chlorophyll? Cotyledons two, carnose. Germination phanerocotylar.

Phytochemistry Hydroxyproline betaines, benzylglucosinolate and myrosinase present. Ellagic acid, proanthocyanidins, alkaloids, and cyanogenic compounds not found. Tannins?

Vegetative anatomy Phellogen? Secondary lateral growth? Vessel elements with simple? perforation plates; lateral pits alternate?, simple? or bordered? pits. Vestured pits? Imperforate tracheary xylem elements ? with bordered? pits, non-septate? Wood rays? Axial parenchyma? Sieve tube plastids S type? Nodes? Myrosin cells? Crystals?

Leaves Opposite, palmately ternate compound (trifoliolate), with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection? Venation palmate. Stomata ?-cytic. Cuticular wax crystalloids? Leaflet margins entire.

Flowers Actinomorphic. Hypogyny. Sepals five to eight, spiral, with imbricate? aestivation, membranous, connate into tube. Petals five to eight, with proximally valvate and distally imbricate aestivation. Androgynophore present. Nectary conical, ascending from petal base to staminal base, surrounding androgynophore. Disc absent.

Androecium Stamens c. 60 to c. 70. Filaments free from each other and from tepals. Anthers dorsifixed?, versatile?, tetrasporangiate, latrorse?, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads?, ?-cellular at dispersal. Exine tectate, with columellate? infratectum, perforate.

Gynoecium Pistil composed of four to six connate carpels. Ovary superior, quadrilocular to sexalocular, stipitate (androgynophore). Style single, simple. Stigma capitate?, type? Pistillodium absent.

Ovules Placentation axile, with ovules in two rows. Ovules several per carpel, bitegmic?, crassinucellar? Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous?, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A many-seeded loculicidal capsule, dehiscing along ventral suture from base to apex, with persistent 4–6-ridged axis.

Seeds Aril? Testa? Tegmen? Perisperm not developed? Endosperm sparse or absent? Embryo curved, poorly differentiated, chlorophyll? Cotyledons two. Germination?

Capparaceae Juss., Gen. Plant.: 242. 4 Aug 1789 [‘Capparides’], nom. cons.; Cruciferae Juss., Gen. Plant.: 237. 4 Aug 1789, nom. cons. et nom. alt.; Drabaceae Martinov, Tekhno-Bot. Slovar: 215. 3 Aug 1820; Erysimaceae Martinov, Tekhno-Bot. Slovar: 238. 3 Aug 1820 [‘Erysimoides’]; Sisymbriaceae Martinov, Tekhno-Bot. Slovar: 583. 3 Aug 1820; Thlaspiaceae Martinov, Tekhno-Bot. Slovar: 633. 3 Aug 1820 [‘Thlaspiceae’]; Cleomaceae Bercht. et J. Presl., Přir. Rostlin 2(64): 253. 1825 [‘Cleomeae’]; Stanleyaceae Nutt. in J. Acad. Nat. Sci. Philadelphia 7: 85. 28 Oct 1834 [‘Stanleae’]; Arabidaceae Döll, Rhein. Fl.: 573. 24-27 Mai 1843 [‘Arabideae’]; Raphanaceae Horan., Char. Ess. Fam.: 169. 30 Jun 1847 [‘Cruciferae s. Raphanaceae’]; Schizopetalaceae A. Juss. in V. V. D. d’Orbigny, Dict. Univ. Hist. Nat. 11: 419. 9 Sep 1848 [‘Schizopetaleae’]; Isatidaceae Döll, Fl. Baden 3: 1310. 30-31 Dec 1861; Oxystylidaceae Hutch., Evol. Phylog. Fl. Pl.: 516. 28 Aug 1969

Fossils Fossil fruits and seeds of Brassicaceae have been described from Oligocene onwards. Fossil pollen grains from the Late Cretaceous of New Zealand have been assigned to Brassicaceae.

Habit Usually bisexual (rarely monoecious, andromonoecious or dioecious), usually perennial, biennial or annual herbs (sometimes evergreen or deciduous shrubs, suffrutices, rarely lianas or small trees). Some species are succulents and some are aquatic. Many species are xerophytes.

Vegetative anatomy Mycorrhiza usually absent. Root hairs absent in Capparoideae and Cleomoideae. Phellogen ab initio usually deeply (cortical etc.; rarely superficially) seated. Cortical and medullary vascular bundles usually absent. Secondary lateral growth usually normal (sometimes anomalous from concentric cambia, e.g., in Boscia) or absent. Vessel elements with simple perforation plates; lateral pits usually alternate (rarely opposite or scattered), bordered pits. Vestured (also on lateral vessel walls) and non-vestured pits present. Imperforate tracheary xylem elements usually fibre tracheids (sometimes libriform fibres) with simple or bordered pits, non-septate (also vasicentric tracheids). Wood rays uniseriate or multiseriate, usually homocellular (sometimes heterocellular). Axial parenchyma apotracheal diffuse, or paratracheal scanty vasicentric, confluent, or banded. Wood elements often storied. Secondary phloem in Capparoideae stratified into hard fibrous and soft parenchymatous layers. Intraxylary phloem present in some genera. Sieve tube plastids usually S type (rarely Pcs type). Endodermis sometimes prominent. Nodes 1:1–3 or 3:3, unilacunar or trilacunar (rarely multilacunar), with one or three leaf traces. Myrosin cells idioblastic. Calciumcarbonate, calciumsulphate and calciumoxalate crystals (prismatic crystals, sometimes druses) present in many representatives (in Capparis also silica).

Trichomes Hairs unicellular or multicellular, uniseriate or multiseriate, simple, furcate or many-branched (occasionally T-shaped malpighiacean hairs), stalked or sessile, medifixed (with two branches parallel to epidermis), stellate (with several or many branches, from common point, parallel to epidermis), candelabra-shaped, dendritic, peltate or lepidote (rarely prickles), often with calciumcarbonate (usually as calcite), or absent; glandular hairs unicellular or multicellular (in Cleomoideae peltate) or absent.

Leaves Alternate (usually spiral, rarely distichous; rarely opposite), usually simple (rarely pinnately or palmately compound), entire or (usually pinnately) lobed, sometimes coriaceous (rarely succulent), with conduplicate ptyxis. Stipules usually absent (rarely present, sometimes replaced by glands or extrafloral nectaries; in Capparis and Cleome sometimes as spines); leaf sheath absent. Petiole vascular bundle transection annular or arcuate. Venation usually pinnate (sometimes palmate, in Pringlea flabellate). Stomata usually anomocytic or anisocytic (sometimes helicocytic, rarely cyclocytic, tetracytic, staurocytic, etc.). Cuticular wax crystalloids as transversely ridged rodlets, chemically dominated by ketones etc. Stomatal myrosin cells absent. Mesophyll cells often with sclerenchymatous idioblasts (with sclereids of different types, e.g., dendrosclereids in Boscia). Epidermis usually with mucilaginous idioblasts; epidermal cells sometimes crystalliferous. Leaf margin serrate, lobate or entire. Extrafloral nectaries rarely present on lamina (e.g. in Crataeva).

Inflorescence Terminal or axillary, raceme, corymb, often branched, spike or umbel-like, or solitary axillary. Bracts and floral prophylls (bracteoles) usually suppressed.

Flowers Usually bisymmetric (sometimes zygomorphic or actinomorphic), often small. Usually hypogyny (rarely half epigyny). Receptacle flat, wide or elongate (sometimes laciniate at apex or with corona), in zygomorphic flowers often with nectariferous appendage or gland inside adaxial sepal; often prolonged into short or long androgynophore or gynophore. Sepals usually 2+2, decussate (rarely three, five or six), usually with imbricate (sometimes valvate or open) aestivation, in one or two whorls, usually caducous, usually free (rarely connate at base; at anthesis rarely forming caducous calyptra). Petals (two to) four (to six), usually with imbricate (rarely contorted or open) aestivation, alternisepalous, of equal size or one pair of larger petals and one pair of smaller petals, usually clawed, free (rarely absent). Nectariferous disc usually extrastaminal (sometimes intrastaminal), annular (sometimes with three or four small episepalous appendages), or absent; nectariferous glands of various shape and insertion (on disc, perianth or stamen).

Androecium Stamens usually 2+4 (rarely one, two, four or up to c. 250), usually haplostemonous or diplostemonous, two outer stamens usually shorter than four inner stamens (tetradynamous), centrifugally developing from four primordia. Filaments filiform (sometimes with appendages), usually in one whorl, usually free (two median ones rarely connate), articulated, free from tepals. Anthers basifixed or dorsifixed, sometimes versatile, tetrasporangiate, introrse or extrorse, longicidal (dehiscing by longitudinal slits); connective sometimes slightly prolonged. Tapetum secretory, with binucleate (to quadrinucleate) cells. Staminodia (four to approx. ten, median-dorsal) present in some zygomorphic species.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (sometimes tricolporate, tricolporoidate, 2–11-colpate, 2–11-colporate or inaperturate), shed as monads, usually tricellular at dispersal (in Capparoideae often bicellular). Exine tectate or semitectate, with columellate infratectum, perforate or reticulate (sometimes smooth, spinulate or finely punctate).

Gynoecium Pistil composed of two (to eight to twelve?) paracarpously connate carpels. Ovary usually superior (rarely semi-inferior), first unilocular, later usually bilocular (in Capparoideae and Cleomoideae sometimes multilocular) with thin secondary septum, usually sessile (in Capparoideae and Cleomoideae usually stipitate, with long gynophore). Style single, usually simple (rarely bifid), often persistent, or absent. Stigma usually one, punctate or capitate (occasionally somewhat lobate), usually papillate, Dry type. Male flowers sometimes with pistillodium?

Ovules Placentation parietal (sometimes intrusively). Ovules one to more than 300 per carpel, usually anatropous or campylotropous (through intrusion of chalazal vascular bundle; rarely amphitropous), usually pendulous or horizontal, bitegmic, usually crassinucellar (rarely tenuinucellar). Micropyle usually bistomal (Z-shaped; rarely endostomal). Outer integument two or three cell layers thick. Inner integument one to four cell layers thick. Endothelium (integumental tapetum) present. Hypostase often present. Megagametophyte monosporous, Polygonum type. Synergids sometimes with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustorium chalazal. Embryogenesis usually onagrad (rarely solanad). Suspensor of various shape, filamentous. Polyembryony occurs.

Fruit A usually septicidal (rarely also loculicidal) capsule, siliqua, dehiscing from base upwards by usually two valves, usually with membranous secondary commissural septum and persistent replum (absent in Capparoideae) consisting of placental tissue (sometimes a nut; rarely a berry, pumpkin fruit, drupe, samara, or schizocarp, lomentum).

Seeds Aril usually absent (present in some Capparoideae). Oily elaiosome present in some species. Seed coat usually endotestal (in Capparoideae and Cleomoideae usually exotegmic, rarely also mesotegmic). Testa non-multiplicative, often winged. Exotesta often palisade. Outer testal layers often mucilaginous. Endotestal cell walls often U-shaped, inner walls thickened, and unthickened wall facing periphery of seed. Tegmen multiplicative. Exotegmic cell walls often (rarely also mesotegmic cells walls) sclerified, fibrous (Capparoideae, Cleomoideae) or non-fibrous (Brassicoideae); exo- and mesotegmen degenerating. Endotegmic cell walls often lignified, often tanniniferous. Perisperm developed in Cadaba, Capparis, Cleome, and Crateva. Endosperm one to more than two cell layers thick or absent. Embryo usually curved or plicate (rarely straight), well differentiated, oily, with chlorophyll. Cotyledons two, flattened or plicate, oily. Radicula dorsal or lateral, usually present in testal pouch. Germination usually phanerocotylar (sometimes cryptocotylar).

Cytology n = (7–)10(–80) (Capparoideae); n = (9–)10(–19)20(–35) (Cleomoideae); n = 4–128; x = (4–)8(–13) (Brassicoideae+Atamisquea: n = 8) – Polyploidy and aneuploidy frequently occurring. Agamospermy present in, e.g., Boechera. Endoplasmic reticulum with protein-rich, wide, organelle-like cisternae.

DNA Plastid gene ndhF with insertion of 6 bp in at least some Capparoideae. Plastid gene infA transferred to nucleus (Arabidopsis). Ancestral KCS6/5 (β-ketoacyl coenzyme A synthases) gene duplicated.

Phytochemistry Flavonols (kaempferol, quercetin; Capparoideae), anthocyanins, ellagic acid, proanthocyanidins (Capparoideae), pyrrolidine alkaloids (Capparoideae), cyanogenic compounds, glucosinolates (synthesized from phenylalanine and/or tyrosine and methionin, chain-extended), methyl glucosinolates (glucocapparine and glucocleomine; Capparoideae and Cleomoideae), sinapic acid, sinapine, cochlearine etc. (choline esters of sinapic acid; seeds of Brassicoideae), ferulic acid, erucic acid, n-eicose-11-enoic acid (Brassicoideae), docosadienoic acid (Brassicoideae), erucic acid, quaternary ammonium compounds (i.a. betaines; Capparoideae and Cleomoideae), and phytoalexins (Brassicoideae) present. Tannins almost absent. Heavy metals accumulated in some species.

Use Ornamental plants, vegetables (Brassica spp., Lepidium sativum, Nasturtium spp.), spices (Brassica spp., Capparis spinosa, Armoracia rusticana, Eutrema wasabi), seed oils (Brassica, Raphanus, Sinapis), forage plants, dyeing substances (Isatis tinctoria), genetic research (Arabidopsis thaliana model plant).

Systematics Brassicaceae is part of a polytomy also comprising Pentadiplandraceae to Tovariaceae.

The taxonomy of Brassicoideae below follows Al-Shehbaz & al. (2006), German & al. (2009) and Warwick & al. (2010).

Neothorelia (1; N. laotica; Thailand, northern Laos) has six sepals and six petals, androgynophore, and very small pollen grains. Its position is unclear, although it may belong somewhere within Brassicaceae or Resedaceae. Neothorelia laotica has sometimes been assigned to Capparoideae (Capparaceae; Williams & Chayamarit 2005).

Capparoideae Burnett, Outlines Bot.: 867. Feb 1835 [‘Capparidae’]

12?/430–440. ‘Crateva’ (10; tropical and subtropical regions; paraphyletic); ‘Capparis’ (c 250; tropical and subtropical regions; diphyletic), Boscia (c 30; tropical and southern Africa, Madagascar, the Arabian Peninsula), Maerua (c 75; tropical regions in the Old World), Bachmannia (1; B. woodii; Mozambique to KwaZulu-Natal and Eastern Cape), Ritchiea (c 30; tropical Africa), Buchholzia (2; B. coriacea, B. tholloniana; tropical West Africa), Euadenia (3; E. brevipetala, E. eminens, E. trifoliolata; tropical Africa), Cladostemon (1; C. kirkii; Mozambique, Zimbabwe, Swaziland, KwaZulu-Natal), Dhofaria (1; D. macleishii; Oman), Cadaba (c 30; southern Africa, Madagascar, Indian Ocean islands, the Arabian Peninsula, India, Australia), Apophyllum (1; A. anomalum; eastern Australia; in Brassicoideae?). – Tropical and subtropical regions. Usually trees or shrubs (sometimes herbs or lianas). Rhizoids (root hairs) absent. Petiole vascular bundle transection annular or arcuate. Leaf base in Crateva glanduliferous. Sclereids present. Inflorescence usually raceme (sometimes fascicle). Flowers actinomorphic or zygomorphic. Sepals usually free (sometimes connate; sepals and petals sometimes fused into tube). Petals four to numerous (sometimes absent). Stamens (one to) four to numerous, centrifugally developing. Filaments usually long. Tectum with various sculpturing. Pistil composed of two to twelve connate carpels; when two carpels, then transversely orientated (superposed-oblique). Gynophore long. Locules sometimes divided by secondary septa. Style usually absent. Outer integument approx. two cell layers thick. Inner integument three or four cell layers thick. Fruit usually indehiscent, baccate (rarely a transversely dehiscent schizocarp or a septicidal capsule). Endotesta sometimes crystalliferous. Tegmen multiplicative, up to six cell layers thick. Exotegmen radially enlarged, sclerified. Anticlinal endotegmic cell walls with lignified bands. n = (7–)10(to more than 15). Insertion of 6 bp in plastid gene ndhF. Methyl glucosinolates and pyrrolidine alkaloids present. – ‘Crateva’ is probably sister group to the remaining Capparoideae. ‘Capparis’ seems to consist of one American clade and one Old World clade, not immediately related to one another.

[Cleomoideae+Brassicoideae]

Usually herbaceous, often annuals. Inflorescence corymb. Stamens six. Pistil composed of two connate carpels. Septicidal capsule with persistent placental vascular strands. Placenta lignified. Seeds 0,5–4 mm long.

Cleomoideae Burnett, Outlines Bot.: 867. Feb 1835 [‘Cleomidae’]

6/260–270. Haptocarpum (1; H. bahiense; eastern Brazil), Cleomella (c 10; southwestern Canada, United States, Mexico), Isomeris (1; I. arborea; California, Mexico), Wislizenia (1–3; W. californica, W. palmeri, W. refracta; southwestern United States, Baja California), Oxystylis (1; O. lutea; Death Valley in southeastern California), Cleome (c 250; tropical and subtropical regions on both hemispheres). – Tropical, subtropical and warm-temperate, with their largest diversity in America. Usually bisexual (some species of Cleome monoecious, with basal male flowers) herbs (Isomeris shrub). C₄ photosynthesis rare (some species of Cleome). Rhizoids (root hairs) absent. Petiole vascular bundle transection arcuate. Leaves usually palmately compound (occasionally simple, palmately lobed). Stipules usually absent. Inflorescence racemose (corymb?). Bracts usually foliaceous. Flowers zygomorphic (sometimes initially bisymmetric). Sepals four, with abaxial sepal often much larger than and in bud covering remaining floral parts. Petals four, clawed. Stamens six. Filaments long. Anthers spirally twisted at dehiscence. Tectum with various sculpturing, often spinulate. Pistil usually composed of two (sometimes four, orthogonally orientated) connate carpels. Gynophore usually present (androgynophore sometimes present). Placenta usually thin, lignified, loop-shaped, persistent in fruit. Outer integument two or three cell layers thick. Inner integument two to ten cell layers thick. Parietal tissue three to five cell layers thick. Nucellar cap approx. two cell layers thick. Endothelium usually absent. Fruit usually capsular (in the Dipterygium group of Cleome a winged single-seeded nut). Seeds usually without aril. Exotegmic cells radially enlarged, sclerified. Endotegmic periclinal cell walls with lignified bands. Suspensor sometimes massive, haustorial. Cotyledons usually incumbent. n = more than 9. Genome duplication present (perhaps c. 20 million years old). Methyl glucosinolates present.

Cladogram of Cleomoideae and Capparoideae based on DNA sequence data (Hall & al. 2002).

Cladogram (simplified) of Cleomoideae pro parte based on DNA sequence data (Riser & al. 2013).

Brassicoideae Prantl, Lehrbuch Bot.: 255. 1 Mar-15 Apr 1880 [‘Brassiceae’]

322–324/2.515–2.645. Cosmopolitan, with their largest diversity in temperate regions in the Northern Hemisphere. Usually herbs (sometimes shrubs, rarely trees). Mycorrhiza absent (vesicular-arbuscular mycorrhiza possibly inhibited by glucosinolates; arbuscular mycorrhiza reported from Thlaspi). Phellogen at least usually deeply seated. Intraxylary phloem rare. Hairs sometimes furcate, stellate or T-shaped. Stipules absent. Stomata anisocytic. Inflorescence racemose (corymbose). Bracts usually absent (sometimes foliaceous). Flowers usually bisymmetric (rarely zygomorphic), with closed development. Petals rarely lobate or fimbriate. Stamens (two, four or) six (24 in Megacarpaea polyandra), two outer often shorter than four inner (tetradynamous), approximately as long as petals. Lateral nectary lobes positioned outside inner stamens. Pollen grains tricellular at dispersal. Tectum often reticulate. Pistil composed of two connate carpels. Ovary usually with commissural septum (absent in Pringlea). Style usually short (rarely long). Gynophore usually absent. Stigma commissural (possibly indicating a basically quadricarpellate gynoecium with axile placentation). Outer integument two to four (or five) cell layers thick. Inner integument (two or) three to eight (to 15) cell layers thick. Parietal tissue approx. one cell layer thick. Hypostase present. Endothelium present. Archespore sometimes multicellular. Fruit usually a siliqua (often explosively dehiscent) with persistent lignified placental strands and membranous secondary septum, replum (fruit occasionally latiseptate or angustiseptate). Seed plicate, but seed coat not bulging inwards. Testa often mucilaginous, three-layered. Radial exotestal cell walls reticulately thickened. Endotesta palisade, lignified, often with U-shaped thickenings, without crystals. Tegmen often multiplicative, non-persistent. Chalazal endosperm cyst present (possibly involved in transport of metabolites into seed; transfer cells present around cyst). Endosperm one-layered. Embryo plicate (sometimes spirally twisted). Cotyledons sometimes conduplicate-incumbent. Radicula not present in testal pouch. x = 4 (before Ata duplication), 7, 8 (after Ata duplication) (–13). Sporophytic self-incompatibility frequent. Duplication of whole genome (Ata palaeopolyploidization). Duplication of nuclear gene PHYB leading to gene PHYD. Nortropane alkaloids sometimes present. Methyl glucosinolates absent. Nickel or zinc accumulated in many species. Selenium accumulated in, e.g., Stanleya pinnata. –Aethionemateae (Aethionemodae) are sister-group to the remaining Brassicoideae.

Aethionemateae Al-Shehbaz, Beilstein et E. A. Kellogg in Plant Syst. Evol. 259: 110. 19 Jul 2006

1/c 55. Aethionema (c 55; the Mediterranean, southeastern Europe and southwestern Asia to Afghanistan). – Annual or perennial herbs or shrubs. Hairs absent. Petal claw three-veined. Nectaries two, lateral. Gynoecium sessile or almost sessile. Ovules (one or) two to four (to eight) per carpel. Fruit a flattened, winged angustiseptate silicula, often nut-like. Testa sometimes mucilaginous. x = 7, 8, 11, 12, 14, 16, 18, 21, 22, 24, 30. Nortropane alkaloids present. – Aethionema is sister to Brassicodae = the remaining Brassicoideae (Koch & al. 2001, etc.).

Brassicodae V. E. Avet. in Biol. Žurn. Armenii 43: 602. 12 Oct 1990 [‘Brassicidinae’]

321–323/2.460–2.590. Eglandular hairs sometimes branched, stellate or T-shaped. Inflorescence rarely with glandular stipules. Pollen grains covered by tryphine (not Pollenkitt). Genome duplication present. – The phylogenetic relationships among the main clades are in part very uncertain.

Scoliaxoneae Al-Shehbaz & S. I. Warwick in Taxon 60: 1161. 4 Aug 2011

1/1. Scoliaxon (1; S. mexicanus; northeastern Mexico). – Hairs simple or branched, minute. Fruit a globose siliqua. – Scoliaxon is not included in any of the studies by German & al. (2009) or Warwick & al. (2010). In the analyses by Warwick & al. (2011) it was identified as a basal lineage.

[Erysimeae+[Cardamineae+[[[Descurainieae+Yinshanieae]+[Lepidieae+Smelowskieae]]+[Camelineae+[[Boechereae+[Halimolobeae+Oreophytoneae]]+[Turritideae+[[Crucihimalayeae+Physarieae]+Pachycladinae+Microlepidieae+[Dipoma+Hemilophia]+Alyssopsideae]]]]]]]

Erysimeae Dumort., Fl. Belg.: 123. 1827

1/c 180. Erysimum (c 180; Europe, Macaronesia, the Mediterranean, Africa, temperate Asia, North America). – Hairs sometimes malpighiaceous. Cardenolides present.

[Cardamineae+[[[Descurainieae+Yinshanieae]+[Lepidieae+Smelowskieae]]+[Camelineae+[[Boechereae+[Halimolobeae+Oreophytoneae]]+[Turritideae+[[Crucihimalayeae+Physarieae]+Pachycladinae+Microlepidieae+[Dipoma+Hemilophia]+Alyssopsideae]]]]]]

Cardamineae Dumort., Fl. Belg.: 124. 1827

c 12/c 345. Aplanodes (2; A. doidgeana: KwaZulu-Natal; A. sisymbrioides: Eastern Cape, Lesotho), Armoracia (3; A. macrocarpa, A. rusticana, A. sisymbrioides; eastern Europe to Siberia, eastern North America), Barbarea (22; Europe, northern Asia, North America), Cardamine (c 200; cosmopolitan), Iodanthus (1; I. pinnatifidus; eastern United States from Connecticut and Minnesota to Texas and Alabama), Leavenworthia (8; southern and southeastern United States), Nasturtium (c 15; temperate regions on the Northern Hemisphere), Ornithocarpa (2; O. fimbriata, O. torulosa; Mexico), Planodes (2; southern United States, northern Mexico), Rorippa (c 85; nearly cosmopolitan, especially temperate regions), Selenia (5; S. aurea, S. dissecta, S. geniculata, S. grandis, S. jonesii; southern United States, Mexico), Sisymbrella (1; S. aspera; western Mediterranean). – Hairs simple or absent. Cotyledons accumbent. Fruit latiseptate or terete (in Armoracia angustiseptate). x = 8. Plastid trnF pseudogene present.

[[[Descurainieae+Yinshanieae]+[Lepidieae+Smelowskieae]]+[Camelineae+[[Boechereae+[Halimolobeae+Oreophytoneae]]+[Turritideae+[[Crucihimalayeae+Physarieae]+Pachycladinae+Microlepidieae+[Dipoma+ Hemilophia]+Alyssopsideae]]]]]

[[Descurainieae+Yinshanieae]+[Lepidieae+Smelowskieae]]

[Descurainieae+Yinshanieae]

Descurainieae Al-Shehbaz, Beilstein et E. A. Kellogg in Plant Syst. Evol. 259: 111. 19 Jul 2006

6/c 47. Descurainia (c 35; Europe, Macaronesia, the Mediterranean, Asia, North and South America), Robeschia (1; R. schimperi; the Middle East; in Descurainia?), Trichotolinum (1; T. deserticola; Patagonia; in Descurainia?), Hornungia (5; H. alpina, H. aragonensis, H. pauciflora, H. petraea, H. procumbens; Europe, the Mediterranean, one species, H. procumbens, a widespread weed), Ianhedgea (1; I. minutiflora; southwestern Asia to China), Tropidocarpum (4; T. californicum, T. capparideum, T. gracile, T. lanatum; California, Mexico, Chile). – Europe, Macaronesia, the Mediterranean, Asia, North and South America. Hairs usually dendritic (rarely dichotomous). Cotyledons incumbent. x = 6(Hornungia), 7.

Yinshanieae Al-Shehbaz, Warwick, Mummenh. et M. A. Koch in Plant Syst. Evol. 285: 229. Mai 2010

1/7–13. Yinshania (7–13; China, northern Vietnam). – Hairs simple, dichotomous or absent. Fruit a latiseptate or angustiseptate silicula. Cotyledons incumbent. – Yinshania is sister to Descurainieae, according to Warwick & al. (2010).

[Lepidieae+Smelowskieae]

Lepidieae DC. in Mém. Mus. Hist. Nat. 7(1): 240. 20 Apr 1821 [‘Lepidineae’]

3/175–220. Cyphocardamum (1; C. aretioides; Afghanistan), Lepidium (175–220; temperate and subtropical regions), Lithodraba (1; L. mendociensis; Argentina). – Cosmopolitan. Hairs simple or absent. Ovule one per locule. Fruit angustiseptate. Seeds often mucilaginous. Cotyledons diplecolobal. x = 8.

Smelowskieae Al-Shehbaz, Beilstein et E. A. Kellogg in Plant Syst. Evol. 259: 111. 19 Jul 2006

1/25. Smelowskia (25; Pakistan to Central and northeastern Asia, northwestern North America). – Hairs branched. Seeds not mucilaginous. Cotyledons incumbent. x = 6 (also n = 10–12). – Smelowskia is sister to Lepidieae.

[Camelineae+[[Boechereae+[Halimolobeae+Oreophytoneae]]+[Turritideae+[[Crucihimalayeae+Physarieae]+Pachycladinae+Microlepidieae+[Dipoma+Hemilophia]+Alyssopsideae]]]]

Camelineae DC. in Mém. Mus. Hist. Nat. 7(1): 239. 20 Apr 1821

8/36. Arabidopsis (11; Europe, tropical African mountains, temperate Asia, North America), Camelina (8; central and southeastern Europe, eastern Mediterranean to central Asia), Capsella (8; eastern Mediterranean, western Asia), Catolobus (1; C. pendulus; eastern Europe, temperate Asia), Chrysochamela (4; C. draboides, C. elliptica, C. noeana, C. velutina; eastern Mediterranean to Russia), Neslia (1; N. paniculata; southeastern Europe, the Mediterranean, southwestern Asia), Noccidium (2; N. hastulatum, N. tuberculatum; southwestern Asia), Pseudoarabidopsis (1; P. toxophylla; Central Asia to western China). – Europe, temperate Asia, northern North America. Hairs simple, stalked or sessile, or stellate. Fruit usually a terete, latiseptate or quadrangular (in Capsella angustiseptate) siliqua (in Camelina, Capsella and Neslia a silicula). x = (6–)8(–11) (x = 4 in Stenopetalum; x = 5 in Arabidopsis thaliana). – Camelineae are probably non-monophyletic.

[[Boechereae+[Halimolobeae+Oreophytoneae]]+[Turritideae+[[Crucihimalayeae+Physarieae]+Pachycladinae+Microlepidieae+[Dipoma+Hemilophia]+Alyssopsideae]]]

[Boechereae+[Halimolobeae+Oreophytoneae]]

Boechereae Al-Shehbaz, Beilstein et E. A. Kellogg in Plant Syst. Evol. 259: 111. 19 Jul 2006

8/c 118. Anelsonia (1; A. eurycarpa; western United States), Boechera (c 110; the Russian Far East, Canada, Greenland, the United States, northern Mexico), Borodinia (1; B. macrophylla; eastern Siberia), Cusickiella (2; C. douglasii, C. quadricostata; western United States), Nevada (1; N. holmgrenii; Nevada), Phoenicaulis (1; P. cheiranthoides; western North America), Polyctenium (1; P. fremontii; western North America), Sandbergia (1; S. whitedii; northwestern United States). – The Russian Far East, North America. Hairs usually branched (in Nevada simple; in Boechera few or absent). x = 7.

[Halimolobeae+Oreophytoneae]

Halimolobeae Al-Shehbaz, Beilstein et E. A. Kellogg in Plant Syst. Evol. 259: 111. 19 Jul 2006

5/45. Exhalimolobos (10; Mexico to South America), Halimolobos (10; southwestern United States to Central America), Mancoa (9; Central and South America), Pennellia (10; southwestern United States, Mexico, Guatemala, Bolivia, northern Argentina), Sphaerocardamum (6; S. compressum, S. divaricatum, S. macropetalum, S. macrum, S. nesliiforme, S. stellatum; Mexico). – Southwestern United States to South America. Hairs branched. Seeds mucilaginous as wet. x = 8. Plastid trnF pseudogene present.

Oreophytoneae Al-Shehbaz, Warwick, Mummenh. et M. A. Koch in Plant Syst. Evol. 285: 229. Mai 2010

2/6. Murbeckiella (5; M. boryi, M. huetii, M. pinnatifida, M. sousae, M. zanonii; southwestern Europe, western Mediterranean, the Caucasus), Oreophyton (1; O. falcatum; tropical mountains in East and Northeast Africa). – Southwestern Europe, northwestern Africa, tropical East Africa. Hairs simple or multifurcate, stipitate, or absent. Fruit a latiseptate siliqua. Cotyledons incumbent.

[Turritideae+[[Crucihimalayeae+Physarieae]+Pachycladinae+Microlepidieae+[Dipoma+Hemilophia]+Alyssopsideae]]

Turritideae Buchenau, Fl. Nordwestdeut. Tiefebene: 258. 28 Apr 1894 [‘Turritinae’]

1/4. Turritis (4; T. brassica, T. chilensis, T. glabra, T. laxa; Europe, Africa, western Asia; T. brassica: mountains in central and southern Europe). – Fruit a siliqua, tetrangular in cross-section. Hairs simple or stellate.

[[Crucihimalayeae+Physarieae]+Microlepidieae+[Dipoma+Hemilophia]+Alyssopsideae]

[Crucihimalayeae+Physarieae]

Crucihimalayeae D. German et Al-Shehbaz in Nord. J. Bot. 28: 647. 15 Dec 2010

3/14. Crucihimalaya (12; Central Asia, the Himalayas, Mongolia), Ladakiella (1; L. klimesii; Ladak in northwestern India), Transberingia (1; T. bursifolia; eastern Russia, western North America, southwestern Greenland). – Northeastern Asia, one species also in North America and southwestern Greenland.

Physarieae B. L. Rob., Syn. Fl. N. Amer. 1: 100. 1895

7/133–138. Dimorphocarpa (4; D. candicans, D. membranacea, D. pinnatifida, D. wislizeni; southwestern Canada, western United States), Dithyrea (2; D. californica, D. maritima; southwestern United States, northwestern Mexico), Lyrocarpa (3; L. coulteri, L. linearifolia, L. xantii; California, Mexico), Nerisyrenia (9; southern United States, Mexico), Paysonia (8; southeastern United States), Physaria (105–110; southwestern Canada, western United States, northwestern Mexico), Synthlipsis (2; S. densiflora, S. greggii; southern United States, Mexico). – Southwestern Canada to northern Mexico, Bolivia, Argentina. Hairs usually sessile, stellate (in Paysonia also simple, dichotomous and stalked substellate). Pollen grains with at least four colpi. Ovules two or more per locule. Fruit an angustiseptate or inflated silicula (in Nerisyrenia sometimes a siliqua). x = 8 (n = 4–11).

Microlepidieae Al-Shehbaz, Warwick, Mummenh. et M. A. Koch in Plant Syst. Evol. 285: 228. Mai 2010 (incl. Pachycladinae O. E. Schulz in Engler et Prantl, Nat. Pflanzenfam. 86: 19, 181. 22 Jul 1924)

16/55. Pachycladon (10; the Southern Alps in New Zealand, Tasmania), Menkea (6; M. australis, M. crassa, M. draboides, M. lutea, M. sphaerocarpa, M. villosula; southern Australia), Cuphonotus (2; C. andraeanus, C. humistratus; arid regions in Australia), Irenepharsus (3; I. magicus, I. phasmatodes, I. trypherus; southeasternmost South Australia, southeastern Victoria and New South Wales), ‘Arabidella’ (7; A. chrysodema, A. eremigena, A. filifolia, A. glaucescens, A. nasturtium, A. procumbens, A. trisecta; drier regions in Australia; polyphyletic), Phlegmatospermum (4; P. cochlearinum, P. drummondii, P. eremaeum, P. richardsii; drier regions in southern Australia), Harmsiodoxa (3; H. blennodioides, H. brevipes, H. puberula; arid regions in Australia), Stenopetalum (10; Australia), Drabastrum (1; D. alpestre; eastern Victoria, southeastern New South Wales), Pachymitus (1; P. cardaminoides; southeastern South Australia, western Victoria, southern New South Wales), Ballantinia (1; B. antipoda; south central Victoria, Tasmania), Geococcus (1; G. pusillus; southern semi-arid regions in Australia), Carinavalva (1; C. glauca; central Australia), Microlepidium (2; M. alatum, M. pilosulum; southwestern Western Australia, coastal South Australia), Scambopus (1; S. curvipes; South Australia), Blennodia (2; B. canescens, B. pterosperma; central Australia). – Australia, New Zealand. Hairs simple or bi- to multifurcate, or sometimes stellate or absent. Fruit an angustiseptate silicula or a terete siliqua. Cotyledons incumbent.

Dipoma clade

1 or 2/1 or 6. Dipoma (1; D. iberideum; southwestern China); Hemilophia (5; H. franchetii, H. pulchella, H. rockii, H. serpens, H. sessilifolia; southwestern China). – Southwestern China. Hairs simple or more or less malpighiaceous (sometimes slightly dichotomous). Fruit a terete or slightly angustiseptate siliqua (Dipoma) or a silicula (Hemilophia). – Dipoma fell into a tetrachotomy also consisting of Physarieae, Microlepidieae and Alyssopsideae, according to Warwick & al. (2010). Hemilophia may possibly belong in this clade.

Alyssopsideae Al-Shehbaz, Warwick, Mummenh. et M. A. Koch in Plant Syst. Evol. 285: 228. Mai 2010

4/9. Alyssopsis (2; A. mollis, A. trinervis; Iran to Central Asia), Calymmatium (2; C. draboides, C. notorrhizum;Central Asia), Dielsiocharis (2; D. bactriana, D. kotschyi; Iran, Tajikistan), Olimarabidopsis (3; O. cabulica, O. pumila, O. umbrosa; eastern Mediterranean to western China). – East Mediterranean to China. Hairs bi- or multifurcate, stalked. Cotyledons accumbent or incumbent.

Alysseae DC. in Mém. Mus. Hist. Nat. 7(1): 231. 20 Apr 1821 [‘Alyssineae’]

24/150–220. Alyssum (100–170; Europe, the Mediterranean and North Africa to the Middle East, western, Central and East Asia; non-monophyletic?), Hormathophylla (1; H. halimifolia; western Mediterranean), Lutzia (1; L. fruticosa; Crete, Karpathos, Astipalea, Kasos), Irania (5; I. compacta, I. membranacea, I. multicaulis, I. pendula, I. umbellata; Iraq, Iran, Afghanistan), Clastopus (2; C. erubescens, C. vestitus; Turkey, Iran, Iraq), Physoptychis (3; P. caspica, P. haussknechtii, P. purpurascens; eastern Turkey to northwestern Iran), Pterygostemon (1; P. spathulatus; northwestern China, eastern Kazakhstan), Degenia (1; D. velebitica; Croatia), Acuston (1; A. lunarioides; Crete and the Kikladhes in Greece), Alyssoides (1; A. utriculata; southern and southeastern Europe, Turkey), Resetnikia (1; R. triquetra; coast of Croatia), Brachypus (1; B. asper; eastern Turkey and Armenia to Iraq, Iran and Central Asia), Fibigia (6; F. clypeata, F. eriocarpa, F. lunarioides, F. macrocarpa, F. obovata, F. suffruticosa; eastern Mediterranean and Ukraine to Iran and Afghanistan), Phyllolepidum (2; P. cyclocarpum, P. rupestre; the Apennines, the Balkan Peninsula, Turkey), Bornmuellera (9; the Balkan Peninsula, Turkey), Takhtajaniella (1; T. globosa; Azerbaijan), Cuprella (2; C. antiatlantica, C. homalocarpa; Morocco, Egypt to Turkey, the Middle East, Iran, Pakistan and the Arabian Peninsula), Aurinia (7; A. corymbosa, A. gionae, A. leucadea, A. moreana, A. petraea, A. saxatilis, A. sinuata; Europe, Turkey), Berteroa (5; B. gintlii, B. incana, B. mutabilis, B. obliqua, B. orbiculata; temperate regions in Eurasia), Lepidotrichum (1; L. uechtritzianum; westernmost coast of the Black Sea in Bulgaria and Turkey), Galitzkya (3; G. macrocarpa, G. potaninii, G. spathulata; Central Asia and Russia to northwestern China and Mongolia), Meniocus (4–7; M. aureus, M. hirsutus, M. linifolius, M. serpyllifolius; Spain, northwestern Africa, southeastern Europe, Ukraine and Turkey to Iran, the Middle East, the Arabian Peninsula, Central Asia, Mongolia and China), Clypeola (9; the Mediterranean and East Europe to the Middle East, Iran, Central Asia and the Arabian Peninsula), Odontarrhena (87; South and Southeast Europe, Turkey, the Caucasus to the Middle East, Central Asia and East Asia, one species, O. obovata, in Alaska and western Canada). – Mainly southern and southeastern Europe, northwestern Africa and southwestern Asia. Hairs stellate. Filaments usually with appendages. Siliqua usually latiseptate or terete (rarely angustiseptate). Seeds often winged. x = 8. – Alysseae are plausible sister-group to the remaining Brassicodae with moderate support (parsimony bootstrap value of 79%).

[Megacarpaeeae+[Heliophileae+Notothlaspideae]]

Megacarpaeeae Kamelin ex D. German in Komarovia 6(2): 83. 19 May 2010

2/12. Megacarpaea (9; Europe, western and Central Asia, the Himalayas, western China), Pugionium (3; P. cornutum, P. dolabratum, P. pterocarpum; Russia, Mongolia, northern China). – Eastern Europe to Central Asia. Hairs simple. Fruit in Megacarpaea a deeply bilobate silicula with single-seeded lobes.

[Heliophileae+Notothlaspi]

Heliophileae DC. in Mém. Mus. Hist. Nat. 7(1): 256. 20 Apr 1821

1/c 90. Heliophila (c 90; southern Africa, especially Northern, Western and Eastern Cape). – Hairs simple or absent. Fruit a siliqua. Cotyledons diplecolobal. x = 10.

Notothlaspideae Al-Shehbaz, Warwick, Mummenh. et M. A. Koch in Plant Syst. Evol. 285: 229. Mai 2010

1/2. Notothlaspi (2; N. australe, N. rosulatum; the Southern Alps in New Zealand). – Hairs simple or absent. Fruit a silicle, angustiseptate. Cotyledons incumbent. – Notothlaspi is sister to Heliophila, according to Warwick & al. (2010).

Iberideae Webb et Berthel., Hist. Nat. Iles Canaries 3(2,1): 92. Nov 1837

2/29. Iberis (27; Europe, the Mediterranean, northwestern Africa), Teesdalia (2; T. coronopifolia, T. nudicaulis; Europe, the Mediterranean, southwestern Asia). – Europe, the Mediterranean, northwestern Africa, southwestern Asia. Hairs simple or absent. Flowers often zygomorphic. Fruit usually strongly angustiseptate, with two seeds.

[Chamireae+[Conringieae+[Coluteocarpeae]]

Chamireae Sond. in Abh. Naturwiss. Verein Hamburg 1: 267. 1846

1/1. Chamira (1; C. circaeoides; Western Cape). – Annual herb. – Chamira may be sister to the clade [Conringieae+Noccaeeae].

[Conringieae+Coluteocarpeae]

Conringieae D. A. German et Al-Shehbaz in Harvard Pap. Bot. 13: 169. 30 Jun 2008

2/8. Conringia (6; C. austriaca, C. clavata, C. grandiflora, C. orientalis, C. persica, C. planisiliqua; southern Europe, the Mediterranean to Central Asia), Zuvanda (2; Z. exacoides, Z. meyeri; southwestern Asia). – Europe to Central Asia. Hairs absent. Fruit a siliqua.

Coluteocarpeae V. I. Dorof. In Turczinanowia 7(3): 51. 28 Sep 2004

3/c 125. Coluteocarpus (1; C. reticulatus; mountains in southwestern Asia), Noccaea (c 120; Europe, North Africa, temperate Asia, North America, Mexico, Patagonia), Pseudosempervivum (4; P. amanum, P. aucheri, P. gurulkanii, P. sempervivum; Turkey, Armenia). – Europe, the Mediterranean, temperate Asia, North America, Patagonia. Hairs absent. Fruit angustiseptate. x = 7.

[Thlaspideae+[Eutremeae+[Brassiceae+Bivonaeeae]+Thelypodieae+[Isatideae+[Sisymbrieae+Ochthodieae]]]]]

Thlaspideae DC. in Mém. Mus. Hist. Nat. 7(1): 234. 20 Apr 1821 [‘Thlaspideae’]

12/36. Thlaspi (4; T. arvense, T. ceratocarpum, T. huetii, T. kochianum; temperate regions of the Northern Hemisphere), Mummenhoffia (2; M. alliacea, M. olivieri; central and southern Europe, East African high mountains), Didymophysa (2; D. aucheri, D. fedtschenkoana; Iran to Central Asia and western Himalayas), Peltaria (4; P. alliacea, P. angustifolia, P. emarginata, P. turkmena; eastern Mediterranean to Iran and Central Asia), Pseudocamelina (3; P. aphragmodes, P. campylocarpa, P. glaucophylla; Iran, northern Iraq, northern Pakistan), Peltariopsis (2; P. grossheimii, P. planisiliqua; eastern Turkey, the Caucasus, northern Iran), Graellsia (8; Morocco, Turkey to Pakistan), Parlatoria (1; P. cakiloidea; Iran), Pachyphragma (1; P. macrophyllum; the Caucasus), Pseudovesicaria (1; P. digitata; the Caucasus), ‘Alliaria’ (3; A. auriculata, A. brachycarpa, A. petiolata; Europe, temperate Asia; diphyletic), Lysakia (1; L. rostrata; northern Iran), Sobolewskia (4; S. caucasica, S. clavata, S. sibirica, S. truncata; eastern Mediterranean to the Caucasus). – Europe, the Mediterranean, southwestern and Central Asia, the Himalayas. Hairs simple or absent. Seed coat striate or coarsely reticulate. x = 7.

[Eutremeae+[Brassiceae+Bivonaeeae]+Thelypodieae+[Isatideae+[Sisymbrieae+Ochthodieae]]]]

Eutremeae Al-Shehbaz, Beilstein et E. A. Kellogg in Plant Syst. Evol. 259: 112. 19 Jul 2006

3/34. Chalcanthus (1; C. renifolius; mountains in Iran, Afghanistan and Central Asia), Eutrema (26; Russia, Central Asia, the Himalayas, eastern Tibet, East Asia, Colorado), Pegaeophyton (7; P. angustiseptatum, P. minutum, P. nepalense,P. purii, P. scapiflorum, P. sulphureum, P. watsonii; Central Asia, the Himalayas to western China). – East Europe, temperate Asia, one species also in North America. Hairs simple or absent. Fruit a siliqua. Cotyledons incumbent. x = 7.

[[Brassiceae+Bivonaeeae]+Thelypodieae+[Isatideae+[Sisymbrieae+Ochthodieae]]]

[Brassiceae+Bivonaeeae]

Brassiceae DC. in Mém. Mus. Hist. Nat. 7(1): 242. 20 Apr 1821

c 36/c 275. Nasturtiopsis (1; N. arabica; North Africa to Israel); Ammosperma (2; A. cinerea, A. variabile; North Africa), Brassica (c 165; Europe, the Mediterranean, temperate Asia), Cakile (c 6; C. arabica, C. constricta, C. edentula, C. geniculata, C. lanceolata, C. maritima; coasts in Europe, the Mediterranean, the Arabian Peninsula and North America; incl. Didesmus?, Erucaria?), Didesmus (2; D. aegyptius, D. bipinnatus; eastern Mediterranean; in Cakile?), Erucaria (10; eastern Mediterranean, the Arabian Peninsula, Iran; in Cakile?), Carrichtera (1; C. annua; Macaronesia, the Mediterranean to Iran), Cordylocarpus (1; C. muricatus; Morocco, Algeria), Crambella (1; C. teretifolia; Morocco), Enarthrocarpus (5; E. arcuatus, E. clavatus, E. lyratus, E. pterocarpus, E. strangulatus; eastern Mediterranean, North Africa), Crambe (c 35; Europe, Macaronesia, the Mediterranean, mountains in northern tropical Africa, western Asia), Douepea (2; D. arabica, D. tortuosa; the Arabian Peninsula, Pakistan), Eremophyton (1; E. chevallieri; Morocco, Algeria, Libya), Fezia (1; F. pterocarpa; Morocco), Foleyola (1; F. billotii; western Sahara), Fortuynia (1; F. garcinii; Iran, Afghanistan, Baluchistan), Guiraoa (1; G. arvensis; Spain), Hemicrambe (3; H. fruticosa, H. fruticulosa, H. socotrana; Morocco, Socotra), Henophyton (1; H. deserti; Morocco to Libya), Kremeriella (1; K. cordylocarpus; northwestern Africa), Moricandia (8; the Mediterranean to Pakistan), Morisia (1; M. monanthos; Corsica, Sardinia), Muricaria (1; M. prostrata; Morocco, Libya), Otocarpus (1; O. virgatus; Algeria), Physorhynchus (2; P. brahuicus, P. chamaerapistrum; southern Iran to northwestern India), Pseuderucaria (2; P. clavata, P. teretifolia; Morocco to Israel), Psychine (1; P. stylosa; North Africa), Quezeliantha (1; Q. tibestica; Sahara), Raffenaldia (2; R. platycarpa, R. primuloides; Morocco, Algeria), Rytidocarpus (1; R. moricandioides; Morocco), Savignya (1; S. parviflora; arid regions in North Africa to the Middle East), Schouwia (1; S. purpurea; North Africa, the Arabian Peninsula), Succowia (1; S. balearica; the Canary Islands, western Mediterranean), Vella (>6; V. anremerica, V. bourgaeana, V. lucentina, V. mairei, V. pseudocytisus, V. spinosa; western Mediterranean), Zilla (2; Z. macroptera, Z. spinosa; North Africa to the Arabian Peninsula), Orychophragmus (3; O. limprichtianus, O. violaceus, O. ziguiensis; China). – Europe, the Mediterranean, central and southern Africa, southwestern Asia, the Atlantic coast of North America. Hairs simple or absent. Fruit usually transversely segmented (heteroarthrocarpous; not in Nasturtiopsis). Cotyledons usually conduplicate (not in Nasturtiopsis). – Brassica in the present sense includes Brassica s.str. (c 40; Europe, the Mediterranean, temperate Asia), Ceratocnemum (1; C. rapistroides; Morocco), Coincya (6; C. cheiranthus, C. cintrana, C. johnstonii, C. monensis, C. richeri, C. wrightii; southern Europe, the Mediterranean), Diplotaxis (c 25; Europe, the Mediterranean to northwestern India), Eruca (1; E. vesicaria; the Mediterranean, northeastern Africa), Erucastrum (c 25; Europe, Macaronesia, the Mediterranean), Hirschfeldia (1; H. incana; the Mediterranean), Raphanus (4; R. caudatus, R. confusus, R. indicus, R. raphanistrum; Europe, the Mediterranean to Central Asia), Rapistrum (2; R. perenne, R. rugosum; Europe, the Mediterranean, West Asia), Sinapidendron (4; S. angustifolium, S. frutescens, S. rupestre, S. sempervivifolium; Madeira), Sinapis (6; S. alba, S. allionii, S. arvensis, S. circinata, S. flexuosa, S. pubescens; Europe, the Mediterranean), Trachystoma (3; T. aphanoneurum, T. ballii, T. labasii; Morocco). Orychophragmus comprises herbs with hairs simple or absent, filaments free from each other, fruit a terete or slightly quadrangular silicula, and cotyledons conduplicate.

Bivonaeeae M. A. Koch et Warwick in Taxon 61: 89. 21 Feb 2012

1/1. Bivonaea (1; B. lutea; Sardinia, Sicily, northwestern Africa). – Fruit a silicula with winged valves.

Thelypodieae Prantl in Engler et Prantl, Nat. Pflanzenfam. III, 2: 154. Mar 1891

29/c 260. Chaunanthus (4; C. acuminatus, C. gracielae, C. mexicanus, C. petiolatus; Mexico), Chilocardamum (4; C. castellanosii, C. longistylum, C. onuridifolium, C. patagonicum; southern Argentina), Chlorocrambe (1; C. hastata; Oregon, Utah), Dictyophragmus (3; D. englerianus, D. lactucoides, D. punensis; Peru, Argentina), Dryopetalon (8; southwestern North America), Englerocharis (2; E. pauciflora, E. peruviana; the Andes), Eremodraba (2; E. intricatissima, E. schulzii; Peru, northern Chile), ‘Hesperidanthus’ (5; H. argillaceus, H. barnebyi, H. jaegeri, H. linearifolius, H. suffrutescens; western North America; polyphyletic), Hollermayera (1; H. silvatica; Chile), Ivania (1; I. juncalensis; northern Chile), Mostacillastrum (c 30; Peru, Bolivia, Argentina), Neuontobotrys (13; Chile, Argentina), Parodiodoxa (1; P. chionophila; mountains in northern Argentina), Petroravenia (1; P. eseptata; Argentina), Phlebolobium (1; P. maclovianum; the Falkland Islands), Phravenia (1; P. vireckii; northern Mexico), Polypsecadium (c 15; northern and western South America), Pringlea (1; P. antiscorbutica; Prince Edward Islands, Crozet Islands, Kerguélen Islands, Heard Island), Romanschulzia (14; Central America), Sarcodraba (4; S. andina, S. dusenii, S. karraikensis, S. subterranea; the Andes), Sibara (13; western and southern North America), Stanleya (6–7; S. albescens, S. bipinnata, S. confertiflora, S. lata, S. pinnata, S. tomentosa, S. viridiflora; western North America), Streptanthus (c 55; western and southern North America; paraphyletic?), Thelypodiopsis (15; North America, Mexico, Guatemala), Thelypodium (19; western North America, Central America), Thysanocarpus (5; T. conchuliferus, T. curvipes, T. erectus, T. laciniatus, T. radians; western United States), Warea (4; W. amplexifolia, W. carteri, W. cuneifolia, W. sessilifolia; southeastern United States), Weberbauera (c 20; the Andes in Peru, Bolivia and Argentina), Zuloagocardamum (1; Z. jujuyensis; northwesternmost Argentina). – North, Central and South America, subantarctic islands.

[Isatideae+[Sisymbrieae+Ochthodieae]]

Isatideae DC. in Mém. Mus. Hist. Nat. 7(1): 241. 20 Apr 1821

5/c 34. Chartoloma (1; C. platycarpum; Central Asia), Glastaria (1; G. glastifolia; southwestern Turkey, Syria, Iraq), Isatis (c 30; Europe, the Mediterranean to Central Asia; paraphyletic), Myagrum (1; M. perfoliatum; Central Europe, the Mediterranean to northwestern India), Schimpera (1; S. arabica; eastern Mediterranean to southern Iran). – Europe, Central and Southwest Asia. Hairs simple or absent. Fruit nutlike, one- or two-seeded, often pendent. x = 7.

[Sisymbrieae+Ochthodieae]

Sisymbrieae DC. in Mém. Mus. Hist. Nat. 7(1): 237. 20 Apr 1821 [‘Sisymbreae’]

1/44. Sisymbrium (44; Europe, the Mediterranean, northern and southern Africa, Asia, one species in North America). – Hairs simple or absent. Stigma bilobate. Fruit a terete siliqua. x = 7.

Ochthodieae Bunge in Arb. Naturf. Ver. Riga 1: 170. 1847

2/2. Ochthodium (1; O. aegyptiacum; eastern Mediterranean); Pseudofortuynia (1; P. esfandiarii; Iran). – Eastern Mediterranean, Iran. Fruit a silicula (in Ochthodium lignified and indehiscent). – Ochthodium and Pseudofortuynia form a clade with weak bootstrap support (less than 60%, according to Warwick & al. 2010), which may be sister to Sisymbrium.

[[Buniadeae+Kernereae]+[Asteae+[Schizopetaleae+[Cremolobeae+Eudemeae+Aphragmeae]]]]

[Buniadeae+Kernereae]

Buniadeae DC. in Mém. Mus. Hist. Nat. 7(1): 245. 20 Apr 1821

1/3. Bunias (3; B. cochlearioides, B. erucago, B. orientalis; southern and eastern Europe, the Mediterranean, western Asia). – Hairs simple or branched, glandular, or absent. Fruit an indehiscent silicula.

Kernereae Al-Shehbaz, Warwick, Mummenh. et M. A. Koch in Plant Syst. Evol. 285: 228. Mai 2010

2/2. Kernera (1; K. saxatilis; mountains in central and southern Europe), Rhizobotrya (1; R. alpina; western Dolomites in central Europe). – Central Europe. Hairs simple or dichotomous, stalked, or absent. Fruit a siliqua, terete or latiseptate. Cotyledons accumbent.

[Asteae+[Schizopetaleae+[Cremolobeae+Eudemeae+Aphragmeae]]]

Asteae Al-Shehbaz, Warwick, Mummenh. et M. A. Koch in Plant Syst. Evol. 285: 228. Mai 2010

1/1. Asta (1; A. schaffneri; Mexico). – Hairs absent. Cotyledons incumbent. . – Asta may be closely allied to Kernereae.

[Schizopetaleae+[Cremolobeae+Eudemeae+Aphragmeae]]

Schizopetaleae R. Br. ex Barnéoud in Ann. Mag. Nat. Hist., ser. 1, 16: 68. Jul 1845

2/20. Atacama (1; A. nivea; northern Chile); Mathewsia (9; southern Peru, northern Chile), Schizopetalon (10; the Andes in northern central Chile and northwestern Argentina). – Southern Peru, northern Chile, northwestern Argentina, mainly in the Atacama desert. Hairs usually simple (sometimes branched) or absent. Fruit usually a siliqua. x = (10–)14. – Atacama was recovered as sister to Teesdalia (Iberideae) in the analysis by Toro-Núñoz & al. (2015). – Schizopetaleae may be sister-group to Cremolobeae.

[Cremolobeae+Eudemeae+Aphragmeae]

Cremolobeae R. Br. in D. Denham et B. H. Clapperton, Narr. Travels Africa: 212. Mar-Apr 1826

3/32. Aimara (1; A. rollinsii; northwestern Chile), Cremolobus (7; C. bolivianus, C. chilensis, C. peruvianus, C. rhomboideus, C. stenophyllus, C. subscandens, C. suffruticosus; the Andes in Peru, Chile and Argentina), Menonvillea (24; the Andes in Chile and Argentina). – Peru, Chile, Argentina. Herbs or shrublets. Hairs simple. Fruit a silicula.

Eudemeae Al-Shehbaz, Warwick, Mummenh. et M. A. Koch in Plant Syst. Evol. 285: 228. Mai 2010

9–10/42–43. Alshehbazia (1; A. hauthalii; the Andes in southernmost Chile and southernmost Argentina), Xerodraba (8; southern Argentina), Onuris (6; O. alismatifolia, O. graminifolia, O. hatcheriana, O. papillosa, O. reichei, O. spegazziniana; Chile, Patagonia), Stenodraba (9; the Andes in Chile and Argentina; diphyletic?), Aschersoniodoxa (4; A. cachensis, A. mandoniana, A. pilosa, A. rusbyi; the Andes), Brayopsis (6; B. alpaminae, B. calycina, B. colombiana, B. diapensioides, B. gamosepala, B. monimocalyx; the Andes), Dactylocardamum (1; D. imbricatifolium; Peru), Delpinophytum (1; D. patagonicum; Patagonia), Eudema (6; E. friesii, E. hauthalii, E. incurva, E. nubigena, E. rupestris, E. werdermannii; the Andes). – The Andes (especially Chile), Patagonia. Hairs simple or dichotomous, stalked, or absent. Fruit a siliqua or silicula, latiseptate or angustiseptate. Cotyledons incumbent. – Horwoodia (1; H. dicksoniae; the Arabian Peninsula to Iraq) was recovered with relatively low support as sister to Eutremeae in the analysis by Toro-Núñez & al. (2015).

Aphragmeae D. A. German et Al-Shehbaz in Harvard Pap. Bot. 13: 168. 30 Jun 2008

1/12. Aphragmus (12; the Himalayas, Siberia, the Russian Far East, Alaska, Yukon). – Hairs simple or dichotomous, small. Fruit a siliqua or silicula. x = 7. – Aphragmus may be closely allied to Eudemeae.

[Calepineae+[Subularieae+[[[Arabideae+Stevenieae]+Biscutelleae]+[[Cochlearieae+Asperuginoides]+[Anchonieae+[Euclidieae+[Dontostemoneae+Chorisporeae]+[Hesperideae+Anastaticeae]]]]]]]

Calepineae Horan, Char. Ess. Fam: 169. 17 Jun 1847

3/7. Calepina (1; C. irregularis; southern and southwestern Europe, Macaronesia, the Mediterranean), ‘Goldbachia’ (5; G. ikonnikovii, G. laevigata, G. pendula, G. tetragona, G. verrucosa; southeastern Russia, Southwest to Central Asia; paraphyletic), Leiocarpaea (1; L. cochlearioides; southestern Russia, western Asia). – Southern Europe, the Mediterranean, Southwest and Central Asia. Hairs simple (often papillary) or absent. Fruit a silicula or siliqua (in ‘Goldbachia’ articulated with single-seeded segments; in Spirorhynchus nut-like).

[Subularieae+[[[Arabideae+Stevenieae]+Biscutelleae]+[[Cochlearieae+Asperuginoides]+[Anchonieae+[Euclidieae+[Dontostemoneae+Chorisporeae]+[Hesperideae+Anastaticeae]]]]]]

Subularieae DC. in Mém. Mus. Hist. Nat. 7(1): 257. 20 Apr 1821

3/4. Idahoa (1; I. scapigera; western United States), Petrocallis (1; P. pyrenaica; mountains in southwestern and central Europe), Subularia (2; S. aquatica: temperate regions in Europe, Asia and North America; S. monticola: mountains in tropical East and Central Africa). – Temperate regions on the Northern Hemisphere. Hairs simple or absent. Fruit a latiseptate silicula. – This clade has weak support (parsimony bootstrap value less than 50%), according to Warwick & al. (2010).

[[[Arabideae+Stevenieae]+Biscutelleae]+[[Cochlearieae+Asperuginoides]+[Anchonieae+[Euclidieae+[Dontostemoneae+Chorisporeae]+[Hesperideae+Anastaticeae]]]]]

[[Arabideae+Stevenieae]+Biscutelleae]

[Arabideae+Stevenieae]

Arabideae DC. in Mém. Mus. Hist. Nat. 7(1): 229. 20 Apr 1821

17/490–495. Abdra (2; A. aprica, A. brachycarpa; the United States), ’Arabis’ (c 60; temperate regions on the Northern Hemisphere, tropical African mountains; paraphyletic), Athysanus (1; A. pusillus; western United States), Aubrieta (16; southern Europe to Iran), Draba (c 390; temperate regions on the Northern Hemisphere, the Andes), Drabella (1; D. muralis; Europe to Turkey and the Caucasus, Morocco), Pachyneurum (1; P. grandiflorum; Central Asia), Baimashania (2; B. pulvinata, B. wangii; Yunnan, Qinghai), Pseudodraba (1; P. hystrix; mountains in Afghanistan and Pakistan), Sinoarabis (1; S. setosifolia; Xizang), Arcyosperma (1; A. primulifolium; Nepal), Borodiniopsis (1; B. alaschanica; western China), Botschantzevia (1; B. karatavica; Kazakhstan), Dendroarabis (1; D. fruticulosa; Altai), Parryodes (1; P. axilliflora; Bhutan), Scapiarabis (4; S. ariana, S. karategina, S. popovii, S. saxicola; northern Pakistan, Central Asia, Tajikistan, Afghanistan, Xinjiang), Tomostima (6; western North America, the Andes). – Temperate regions on the Northern Hemisphere, tropical African mountains, the Andes. Hairs branched. Cotyledons usually accumbent (in Berteroella incumbent). Seeds not mucilaginous. x = 8 (polyploidy frequent in Draba).

Stevenieae Al-Shehbaz, D. German et M. A. Koch in Plant Divers. Evol. 129: 72. 22 Mar 2011

3/8. Macropodium (2; M. nivale, M. pterospermum; Central Asia, Sakhalin, Japan), Pseudoturritis (1; P. turrita; southern and southeastern Europe), Stevenia (5; S. alyssoides, S. cheiranthoides, S. incarnata, S. maximowiczii, S. sergievskajae; China, the Korean Peninsula, Japan, the Russian Far East). – Europe, Central and East Asia.

Biscutelleae Dumort., Fl. Belg.: 118. 1827

5/61. Heldreichia (1; H. bupleurifolia; Turkey to Afghanistan), Biscutella (46; central and southern Europe, the Mediterranean), Megadenia (1; M. pygmaea; Siberia, China), Ricotia (9; eastern Mediterranean), Lunaria (4; L. annua, L. elongata, L. rediviva, L. telekiana; Europe). – Europe, the Mediterranean, Asia. Fruit a compressed silicula (in Biscutella didymous, indehiscent); Ricotia has a silique or a latiseptate silicula; Lunaria has a latiseptate silicula and an unusually long gynophore. – Biscutelleae are sister to [Arabideae+Stevenieae].

[[Cochlearieae+Asperuginoides]+[Anchonieae+[Euclidieae+[Dontostemoneae+Chorisporeae]+[Hesperideae+Anastaticeae]]]]

[Cochlearieae+Asperuginoides]

Cochlearieae Buchenau, Fl. Nordwestdeut. Tiefebene: 245. 28 Apr 1894 [‘Cochleariinae’]

2/c 35. Cochlearia (c 30; Western Europe, the western Mediterranean, northwestern Africa, Arctic regions in northeastern Asia, Alaska and northern Canada), Ionopsidium (5; I. abulense, I. acaule, I. albiflorum, I. prolongoi, I. savianum; western Mediterranean). – Europe, the Mediterranean, North Africa, northeastern Asia, Alaska, northern Canada. Hairs absent. Stigma entire. Fruit a terete or angustiseptate silicula. Seeds biseriate. x = 6–7. – Cochlearieae may be sister-group to the remaining Brassicodae. A sister-group relationship between Cochlearieae and Asperuginoides is weakly supported in the analyses by Warwick & al. (2010).

Asperuginoides clade

1/1. Asperuginoides (1; A. axillaris; Turkey and eastwards to Central Asia and Pakistan). – Hairs stellate. Fruit a dorsally compressed silicula, covered with anchor-shaped hairs (glochids). – Asperuginoides may be sister to Cochlearieae, although its position in Brassicaceae is very uncertain.

[Anchonieae+[Euclidieae+[Dontostemoneae+Chorisporeae]+[Hesperideae+Anastaticeae]]]

Anchonieae DC. in Mém. Mus. Hist. Nat. 7(1): 242. 20 Apr 1821

9/66–68. Anchonium (2; A. billardieri, A. elichrysifolium; West and Central Asia), Eremoblastus (1; E. caspicus; western Kazakhstan to Central Asia), Iskandera (2; I. alaica, I. hissarica; Central Asia), ’Matthiola’ (48–50; western Europe, Macaronesia, the Mediterranean; polyphyletic), Micrantha (1; M. multicaulis; Iran), Microstigma (3; M. brachycarpum, M. deflexum, M. sajanense; Siberia, Mongolia, China), Sterigmostemum (6; S. acanthocarpum, S. caspicum, S. incanum, S. longistylum, S. ramosissimum, S. sulphureum; southeastern Russia, western Kazakhstan, Southwest and Central Asia), Synstemon (2; S. lulianlianus, S. petrovii; China), Zerdana (1; Z. anchonioides; mountains in Iran). – Europe, temperate Asia. Hairs branched; glandular hairs multicellular with multiseriate stalk. Stigma often strongly bilobate. x = 7 (x = 5–8 in Matthiola).

[Euclidieae+[Dontostemoneae+Chorisporeae]+[Hesperideae+Anastaticeae]]

Euclidieae DC. in Mém. Mus. Hist. Nat. 7(1): 236. 20 Apr 1821

28/165. Anzhengxia (1; A. yechengnica; Xinjiang Prov. in China), Atelanthera (1; A. perpusilla; Central Asia, the Himalayas), Braya (23; northern circumpolar, mountains in Central and North Europe, Central Asia, the Himalayas), Catenulina (1; C. hedysaroides; Central Asia), Christolea (5; C. borealis, C. crassifolia, C. maidantalica, C. niyaensis, C. parryoides; Central Asia, the Himalayas, East Asia), Cryptospora (3; C. falcata, C. inconspicua, C. trichocarpa; Central Asia), Cymatocarpus (3; C. grossheimii, C. heterophyllus, C. pilosissimus; Transcaucasia to Central Asia), Dichasianthus (1; D. subtilissimus; Central Asia), Dilophia (2; D. ebracteata, D. salsa; Central Asia to western China), Euclidium (1; E. syriacum; eastern Europe to the Middle East), Lachnoloma (1; L. lehmannii; Iran to China), Leiospora (8; Russia to East Asia), Lepidostemon (5; L. everestianus, L. glaricola, L. gouldii, L. pedunculosus, L. rosularis; the Himalayas), Leptaleum (1; L. filifolium; eastern Mediterranean to Central Asia and Baluchistan), Metashangrilaia (1; M. forrestii; Xizang, Yunnan, Bhutan), Neotorularia (13; the Mediterranean to Central Asia and Afghanistan), Octoceras (1; O. lehmannianum; Iran and Afghanistan to Central Asia), Pycnoplinthopsis (1; P. bhutanica; Bhutan), Pycnoplinthus (1; P. uniflora; the Himalayas), Rhammatophyllum (9; Central Asia), Rudolf-kamelinia (1; R. korolkowii; Central Asia, western China, northern India), Shangrilaia (1; S. nana; Yunnan), Sisymbriopsis (5; S. mollipila, S. pamirica, S. schugnana, S. shuanghuica, S. yechengnica; Tajikistan, China), Solms-laubachia (33; China, one species also in Sikkim), Spryginia (7; S. afghanica, S. crassifolia, S. falcata, S. gracilis, S. pilosa, S. undulata, S. winkleri; Central Asia), Streptoloma (2; S. desertorum, S. sumbarense; Central Asia to Afghanistan), Strigosella (24; the Mediterranean to western and Central Asia, North Africa), Tetracme (10; eastern Mediterranean to Central Asia and Baluchistan). – Europe, temperate and arctic Asia, arctic and alpine North America. Hairs usually branched (rarely simple or absent). Multicellular glands absent. Stigma entire or bilobate. Fruit often a terete to quadrangular siliqua or silicula. Cotyledons incumbent. x = 7(–8).

[[Dontostemoneae+Chorisporeae]+[Hesperideae+Anastaticeae]]

[Dontostemoneae+Chorisporeae]

Dontostemoneae Al-Shehbaz et Warwick in Harvard Pap. Bot. 12(2): 431. 21 Dec 2007

2/19. Clausia (6; C. agideliensis, C. aprica, C. kasakhorum, C. podlechii, C. robusta, C. trichosepala; southeastern Europe to Central and East Asia), Dontostemon (13; Siberia, Mongolia, China). – Southeastern Europe to Siberia and East Asia. Perennial herbs. Eglandular hairs simple or dichotomous; glandular hairs when present multicellular, multiseriate. Fruit a terete or latiseptate often tardily dehiscent siliqua. Radicula accumbent.

Chorisporeae Ledeb., C. A. Mey. et Bunge, Fl. Altaic. 3: 104. Jul-Dec 1831

4/57. ’Chorispora’ (13; eastern Mediterranean to Central Asia; paraphyletic), Diptychocarpus (1; D. strictus; eastern Europe to Central Asia), Litwinowia (1; L. tenuisissima; southwestern Asia to China), Parrya (42; northern temperate and arctic regions on the Northern Hemisphere, circumpolar). – Europe, temperate Asia, northern North America. Hairs simple; glandular hairs multicellular with multiseriate stalk. Stigma connivent. Fruit a siliqua or a moniliform schizocarp (lomentum) with one-seeded nutlike mericarps. x = 7 (in Chorispora also x = 9).

[Hesperideae+Anastaticeae]

Hesperideae Prantl in Engler et Prantl, Nat. Pflanzenfam. III, 2: 154. Mar 1891

2/c 30. Hesperis (29–30; Europe, the Mediterranean to Iran, Central Asia and western China), Tchihatchewia (1; T. isatidea; Turkey). – Europe, temperate Asia. Hairs usually simple and/or dichotomous (rarely dendritic); glandular hairs with uniseriate stalk and unicellular head (glands rarely absent). Stigmatic lobes decurrent. Fruit a siliqua. x = 6–10.

Anastaticeae DC. in Mém. Mus. Hist. Nat. 7(1): 236. 20 Apr 1821

13/96. Anastatica (1; A. hierochuntica; Morocco to southern Iran), Cithareloma (2; C. lehmannii, C. vernum; Iran, Central Asia), Diceratella (10; northeastern tropical Africa, Iran), Eigia (1; E. longistyla; Israel to northwestern Arabian Peninsula), Eremobium (1; E. aegyptiacum; North Africa, the Middle East), Farsetia (28; Morocco to northwestern India, mountains in Tanzania), Lachnocapsa (1; L. spathulata; Socotra), Lobularia (5; L. arabica, L. canariensis, L. libyca, L. marginata, L. maritima; Macaronesia, the Mediterranean to the Arabian Peninsula),‘Malcolmia’ (35; the Mediterranean to Afghanistan; polyphyletic), Maresia (3; M. meyeri, M. nana, M. pulchella; the Mediterranean to the Caspian area and southern Iran), Morettia (3; M. canescens, M. parviflora, M. philaeana; North Africa to the Arabian Peninsula), Notoceras (1; N. bicorne; the Canary Islands, the Mediterranean to northwestern India), Parolinia (5; P. filifolia, P. intermedia, P. ornata, P. platypetala, P. schizogynoides; the Canary Islands). – Macaronesia and the Mediterranean to Central Asia and northwestern India and East African mountains. Hairs simple or branched, or absent. Fruit a siliqua or a latiseptate silicula.

Strict consensus tree (simplified) of Brassicoideae based on DNA sequence data (Warwick & al. 2010).

Bayesian inference tree (simplified) of Brassicoideae based on DNA sequence data (German & al. 2009).

Unplaced Brassicoideae

Andrzeiowskia (1; A. cardamine; southeastern Europe, southwestern Asia), Raphanorhyncha (1; R. crassa; Mexico), Veselskya (1; V. griffithiana; Afghanistan).

Papayaceae Blume, Bijdr. 15: 940. Jul-Dec 1826, nom. illeg.; Papayales Blume in C. F. P. von Martius, Consp. Regn. Veg.: 32. Sep-Oct 1835 [‘Papayaceae’], nom. illeg.; Caricales L. D. Benson, Pl. Classif.: 648. 1957

Habit Usually dioecious (sometimes monoecious, andromonoecious, gynomonoecious, or polygamomonoecious; rarely bisexual), evergreen trees or shrubs (rarely perennial herbs or climbing shrubs; in Jarilla and Jacaratia corumbensis with a stout tuberous stem). Stem usually unbranched, often spiny.

Vegetative anatomy Phellogen? Xylem, except vessel elements, unlignified. Vessel elements with simple perforation plates; lateral pits alternate or scalariform, bordered? pits. Non-vestured pits present. Imperforate tracheary xylem elements? Wood rays multiseriate?, heterocellular? Axial parenchyma paratracheal vasicentric, usually non-lignified. Wood elements often storied. Secondary phloem stratified into hard fibrous and soft parenchymatous layers. Phloem fibres as isolated strands. Sieve tube plastids S type. Nodes ≥3:≥3?, trilacunar or multilacunar with three or more? leaf traces. Parenchyma with cellular (multinucleate as mature), articulated, anastomosing laticifers containing latex-like exudate. Idioblastic myrosin cells absent. Calciumoxalate druses present.

Trichomes Eglandular hairs absent; stem and leaves (especially basal adaxial parts of lamina) in Carica with multicellular, uniseriate, simple glandular hairs; leaves in Carica and Jacaratia with adaxial glandular hairs crowned by multicellular head. Stem apex and inflorescence in Carica with colleters.

Leaves Alternate (spiral), usually palmately compound (leaflets sometimes pinnately lobed) or palmately lobed (leaves rarely entire or pinnately lobed), with flat-curved to involute ptyxis. Stipules usually absent (in Vasconcellea stipulata modified into spines); leaf sheath absent. Petiole vascular bundle transection? Venation usually palmate (rarely pinnate). Stomata anomocytic. Cuticular wax crystalloids as rosettes (Fabales type). Lamina with laticifers. Myrosin cells stomatal. Mesophyll with cells containing calciumoxalate druses. Leaf margin serrate or entire.

Flowers Actinomorphic. Hypogyny. Sepals (four or) five, with open aestivation, free or entirely or partially connate. Petals (four or) five, with contorted or valvate aestivation, in male flowers connate into tube, in female flowers free or connate in lower part. Nectaries in Carica and Jacaratia on pistillodium in male flowers; female flowers without nectaries. Disc absent.

Androecium Stamens usually (four or) five longer antesepalous and (four or) five shorter antepetalous, diplostemonous (in Carica sometimes four or five, antesepalous). Filaments sometimes hairy at base (in Carica and Jarilla submoniliform hairs), free or connate at base, adnate to petals (epipetalous). Anthers dorsifixed to subbasifixed, versatile?, usually tetrasporangiate (anthers of inner stamens rarely disporangiate), introrse, longicidal (dehiscing by longitudinal slits); connective often dorsally widened, often with glandular apex. Tapetum secretory. Staminodia in male flowers sometimes (four or) five (absent in female flowers).

Pollen grains Microsporogenesis simultaneous. Pollen grains tricolporate, shed as monads, bicellular at dispersal. Exine tectate or semitectate, with columellate infratectum, perforate or reticulate.

Gynoecium Pistil composed by (four or) five connate carpels. Ovary superior, unilocular (Carica, Horovitzia, Jarilla) or divided by septa and secondarily (quadrilocular or) quinquelocular (Cylicomorpha, Jacaratia, Vasconcellea). Stylodia (four or) five, short, free, or style single, (quadrilobate or) quinquelobate, or absent. Stigmas flabellate or almost petaloid (rarely one capitate-truncate stigma), papillate, Dry type. Female flowers in Jacaratia with white broad stigmas resembling stamens of male flowers. Male flowers often with pistillodium.

Ovules Placentation intrusively parietal (when ovary unilocular) or axile (when ovary multilocular). Ovules usually numerous per carpel, anatropous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument ? cell layers thick. Inner integument four to six cell layers thick. Megagametophyte usually monosporous, Polygonum type (rarely tetrasporous). Synergids with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis via irregular early divisions.

Seeds Aril usually absent (sometimes present). Seed coat testal. Testa multiplicative, with outer epidermis modified into mucilaginous sarcotesta. Mesotestal cell walls with tannins and lignified ridges. Endotesta crystalliferous sclerotesta, sometimes lignified. Tegmen not or only poorly multiplicative. Exotegmen fibrous or non-fibrous, lignified. Endotegmen unspecialized. Perisperm not developed. Endosperm copious, carnose, oily and proteinaceous. Embryo straight, well differentiated, without chlorophyll. Cotyledons two, wide, flattened. Hypocotyl swollen (often tuberous). Epicotyl swollen. Germination phanerocotylar.

Phytochemistry Oleanolic acid derivatives, gallic acid, alkaloids, saponins, benzylglucosinolates synthesized from phenylalanine and/or tyrosine, benzylisothiocyanate (in wax layer of fruit), myo-inositol, and proteolytic enzymes (carpain, papain in latex) present. Cyclopentenoid cyanogenic glycosides? Flavonols, ellagic acid and proanthocyanidins not found.

Use Ornamental plants, fruits, papain for medical and technical purposes (softening of meat, tanning of leather).

Vegetative anatomy Phellogen ab initio cortical. Cambium storied? Vessel elements with simple perforation plates; lateral pits usually alternate (sometimes opposite). Imperforate tracheary xylem elements fibre tracheids? with bordered pits. Wood rays uniseriate or biseriate, homocellular? Axial parenchyma? Sieve tube plastids S type? Nodes? Mesophyll with branched sclereids. Crystals?

Leaves Opposite, simple, entire, with ? ptyxis. Stipules minute; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate? Stomata anomocytic. Cuticular wax crystalloids? Leaf margin entire.

Inflorescence Flowers axillary, solitary. Floral prophylls (bracteoles) absent.

Flowers Zygomorphic, resupinate (twisted 180°). Hypogyny. Sepals five, with ? aestivation, connate into tube in lower half; median sepal abaxial; calyx tube adaxially split to base into two halves. Petals two, with valvate aestivation, latero-abaxial, alternisepalous, sessile?, only with epidermis connate slipper-shaped. Nectary extrastaminal, triangular, inserted on receptacle between petal bases and staminal bases. Receptacle elongated into flattened adaxially curved androgynophore, at apex more or less hidden behind petals.

Androecium Stamens four abaxial fertile, and four or five adaxial staminodial, diplostemonous, forming ring (torus) at apex of androphore; median stamens absent. Filaments connate at base, free from tepals. Anthers basifixed?, non-versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum glandular? Staminodia four or five.

Pollen grains Microsporogenesis simultaneous? Pollen grains usually shortly tricolporate (sometimes tetracolporate) with bulging apertures, shed as monads, tricellular? at dispersal. Exine tectate, with columellate? infratectum, punctate, smooth?

Gynoecium Pistil composed of two or three connate carpels. Ovary superior, bilocular or trilocular (unilocular?), shortly bialate, sessile, on androgynophore. Style absent. Stigma somewhat bilobate or trilobate, type? Pistillodium absent.

Ovules Placentation axile-basal. Ovule one per carpel, ?-tropous, bitegmic?, crassinucellar? Funicle lobed. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type? Endosperm development? Endosperm haustoria? Embryogenesis?

Seeds Aril formed from apex of outer integument. Testa thick (multiplicative?). Exotesta? Endotesta? Tegmen? Perisperm not developed. Endosperm sparse. Embryo curved (conduplicate-involute), well differentiated, chlorophyll? Cotyledons two. Germination?

de Jussieu in V. V. D. d’Orbigny, Dict. Univ. Hist. Nat. 6: 450. 1845 [‘Gyrostemoneae’], nom. cons.

Gyrostemonales Takht., Divers. Classif. Fl. Pl.: 123. 24 Apr 1997; Gyrostemonanae Takht., Divers. Classif. Fl. Pl.: 123. 24 Apr 1997

Distribution Australia, Tasmania, with their highest diversity in southwestern Western Australia.

Habit Usually dioecious (in Walteranthus monoecious), evergreen or deciduous shrubs or small trees (Cypselocarpus and Tersonia annual or biennial herbs). Xerophytes.

Vegetative anatomy Phellogen ab initio subepidermal. Cambium more or less storied. Vessel elements with simple perforation plates; lateral pits alternate or scalariform, simple pits. Non-vestured pits present. Imperforate tracheary xylem elements tracheids with bordered pits, non-septate. Wood rays uniseriate or multiseriate, homocellular. Axial parenchyma, usually paratracheal scanty vasicentric or in tangential bands (sometimes apotracheal diffuse or diffuse-in-aggregates). Wood elements partially storied (shorter imperforate tracheary elements storied). Sieve tube plastids Ss type. Nodes? Myrosin cells idioblastic. Secretory cavities absent. Crystals absent.

Leaves Alternate (spiral), simple, entire, often carnose or coriaceous, with flat ptyxis. Stipules minute or absent; leaf sheath absent. Petiole vascular bundle transection arcuate. Venation pinnate or leaves one-veined. Stomata anomocytic. Cuticular wax crystalloids absent. Stomatal myrosin cells absent. Leaf margin entire.

Inflorescence Terminal or axillary, simple or compound raceme or spike, or flowers solitary axillary.

Flowers Actinomorphic, small. Receptacle flattened, discoid. Hypogyny. Tepals (sepals?) four or five (to eight), in one whorl, with imbricate aestivation, persistent, connate into cup. Petals probably absent. Nectary absent. Disc usually central.

Androecium Stamens seven to more than 100, usually in one whorl inserted on receptacular margin (in some species of Gyrostemon in several whorls covering receptacle), centripetally developing. Filaments very short or absent, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate cells. Female flowers with or without staminodia?

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolpate (sometimes dicolpate or tetracolpate), shed as monads, bicellular at dispersal. Exine tectate, with granular infratectum, psilate to scabrate. Ectexine spongy, undifferentiated.

Gynoecium Pistil composed of usually several to numerous (up to c. 60; in Gyrostemon few; in Cypselocarpus one) free or connate carpels usually in one whorl (sometimes two whorls); when two carpels, then transversely orientated. Ovary superior, unilocular (apocarpy) or several- to multilocular (as many as carpels). Stylodia as many as carpels (sometimes marginal), free or connate in lower part (rarely cleft). Stigmas decurrent, large and often wide, type?; stigmatic ring forming corona. Male flowers sometimes with pistillodium?

Ovules Placentation apical-axile (when syncarpy) or marginal (when apocarpy). Ovule one per carpel, campylotropous (anatropous?), apotropous, bitegmic, crassinucellar. Micropyle bistomal. Outer integument two cell layers thick. Inner integument five cell layers thick. Archespore unicellular (Gyrostemon) or bicellular to quadricellular (Codonocarpus). Nucellar cap absent. Hypostase present. Obturator absent. Megagametophyte monosporous, Polygonum type. Porogamy. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Fruit A dry or carnose schizocarp, with two to numerous ventrally (Gyrostemon) or marginally (Codonocarpus) dehiscing samaroid to follicular mericarps, a syncarp or nutlet (Cypselocarpus, Tersonia, Walteranthus), often with persistent calyx.

Seeds Funicular exostomal aril formed from apex of outer integument, surrounding hilum. Testa not multiplicative. Exotestal cells compressed. Endotestal cells small, cuboid. Tegmen not multiplicative. Exotegmen fibrous. Exotegmic cells elongate, thick-walled. Remaining tegmic cells crushed. Perisperm not developed. Endosperm copious, carnose, oily. Embryo curved, well differentiated, chlorophyll? Cotyledons two. Germination?

Phytochemistry Glucosinolates (synthesized from?) and cyanogenic compounds present. Ellagic acid and proanthocyanidins not found. Tannins?

Engler in Engler et Prantl, Nat. Pflanzenfam., III, 6: 319. 14 Mai 1895, nom. cons.

Habit Bisexual, evergreen shrubs or small trees. Xerophytes. Branches photosynthesizing, terminating in an elongated spine. Prophylls basal on branches and spines.

Vegetative anatomy Phellogen ab initio pericyclic. Cortex with bast bundles and brachysclereids and in older parts secretory resinous ducts. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured and non-vestured pits present. Imperforate tracheary xylem elements tracheids and fibre tracheids with bordered pits, non-septate (also vasicentric tracheids). Wood rays usually multiseriate, homocellular. Axial parenchyma usually apotracheal diffuse or diffuse-in-aggregates, or paratracheal scanty vasicentric. Wood elements not storied. Sieve tube plastids S type? Nodes? Intercellular canals present. Phloem with resinous ducts. Idioblastic myrosin cells present. Crystals? Druses absent.

Leaves Alternate (spiral), simple, entire, very small, scale-like, early caducous, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection? Venation? Stomata sunken. Cuticular wax crystalloids? Idioblastic myrosin cells present. Leaf margin entire.

Flowers Actinomorphic, small and indistinct. Hypogyny. Sepals usually 2+2, decussate (sometimes five), with imbricate aestivation, free. Petals four (or five), diagonal, with imbricate aestivation, shortly clawed, caducous, free. Nectaries inserted at staminal bases. Disc absent.

Androecium Stamens 4+4 (or 5+5), diplostemonous. Filaments flattened, with basal appendages, free from each other and from tepals. Anthers dorsifixed, versatile?, tetrasporangiate, introrse (antesepalous stamens) or latrorse (antepetalous stamens), longicidal (dehiscing by longitudinal slits). Tapetum secretory, with binucleate to multinucleate cells. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colporate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, striate-reticulate.

Gynoecium Pistil composed of two connate carpels, obliquely orientated. Ovary superior, bilocular (or trilocular), shortly stipitate (gynophore present). Style single, simple, subulate, persistent. Stigma slightly bifid, minutely expanded, type? Pistillodium absent.

Ovules Placentation axile. Ovules approx. ten per carpel, campylotropous, apotropous or epitropous, bitegmic, tenuinucellar. Micropyle bistomal, Z-shaped (zig-zag). Outer integument two cell layers thick, non-multiplicative. Inner integument three cell layers thick. Obturator absent. Apical cells of megasporangium much enlarged (functioning as “obturator”?). Hypostase absent. Parietal tissue absent. Megasporangial epidermal cells radially enlarged. Megagametophyte monosporous, Polygonum type. Antipodal cells ephemeral. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Seeds Arilloid small, surrounding hilum (developed from apex of outer integument or exostome). Seed coat exotestal-tegmic. Testa non-multiplicative. Exotesta with massive cuticle and tannin-like substance; exotestal cells large and thick-walled. Endotestal cells tiny. Tegmen non-multiplicative. Exotegmen fibrous, with elongate cells and strongly thickened lignified cell walls. Mesotegmen crushed. Endotegmic cells thick-walled, with tannin-like substance, almost collapsed. Perisperm not developed. Endosperm sparse, thin. Embryo curved, well differentiated, with chlorophyll. Cotyledons two, incumbent. Germination?

Phytochemistry Virtually unknown. Ellagic acid? Tannins? Glucosinolates not found.

Brown in London Edinburgh Philos. Mag. & J. Sci. 3: 70. Jul 1833 [’Limnantheae’], nom. cons.

Limnanthales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 56. Sep-Oct 1835 [’Limnantheae’]; Limnanthineae Engl., Syllabus, ed. 2: 143. Mai 1898

Habit Bisexual, annual herbs. Helophytes, sometimes aquatic. Main root early wilting and replaced by adventitious roots.

Vegetative anatomy Phellogen absent. Primary vascular tissue cylinder of vascular bundles. Secondary lateral growth absent. Wood elements often only in root-stem junction. Vessel elements with simple perforation plates; lateral pits alternate to pseudoscalariform, simple pits. Non-vestured pits present. Xylem without imperforate tracheary elements. Wood rays uniseriate or biseriate, homocellular. Axial parenchyma paratracheal scanty. Wood non-storied. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Myrosin cells idioblastic. Wood without crystals.

Trichomes Hairs unicellular (sometimes verrucose) or absent; glandular hairs absent.

Leaves Alternate (spiral), once or twice pinnately compound, or simple and pinnately lobed, with conduplicate ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Myrosin cells abundant; stomatal myrosin cells absent. Tanniniferous idioblasts present. Leaf margin serrate. Apices of leaf and leaflets sometimes with hydathodes.

Inflorescence Terminal, raceme or flowers solitary axillary. Bracts and floral prophylls (bracteoles) sometimes absent.

Flowers Actinomorphic. Hypogyny or half epigyny. Sepals three or (four or) five, with valvate to somewhat imbricate aestivation, persistent, free or slightly connate at base, comparatively large. Petals three or (four or) five, with dextrorsely contorted aestivation, caducous, free. Nectaries present on swollen abaxial bases of antesepalous (alternipetalous, rarely also antepetalous) stamens, or absent. Disc absent.

Androecium Stamens usually 3+3 or 5+5 (rarely three or 4+4), of two different lengths with longest stamens antesepalous, usually diplostemonous. Filaments free from each other and from tepals. Anthers dorsifixed, versatile, tetrasporangiate, ab initio introrse, subsequently extrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains zonocolporate (syncolpate?) of unique type with two zonocolpi parallel to equator alternatively one colpus parallel to and encircling equator, heteropolar with smaller distal and larger proximal pole, shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, microperforate, striate, spinulate.

Gynoecium Pistil composed of two or three or (four or) five carpels, bulging (seemingly almost free in lower part); when three median carpels abaxial then carpels antesepalous; carpellary walls without vascular bundles. Ovaries superior or partially semi-inferior, two or three or (four or) five, unilocular, free. Style gynobasic, filiform, hollow, bilobate to quinquelobate. Stigmas minutely capitate to punctate, papillate, Dry type. Pistillodium absent.

Ovules Placentation basal-parietal. Ovule one per carpel, anatropous, ascending, apotropous (or epitropous?), unitegmic, tenuinucellar. Integument 14 to 16 cell layers thick. Parietal tissue absent. Archespore multicellular. Megagametophyte tetrasporous, pseudomonosporous (quadrinucleate or sexanucleate with one nucleus developing), modified Drusa type (one or two antipodal cells absent). Synergids sometimes with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria laterally micropylar (on funicular side) or absent. Embryogenesis onagrad. Suspensor uniseriate.

Fruit A schizocarp, with persistent and sometimes accrescent calyx and with (two or) three or five muriculate (spinulate) one-seeded nutlike mericarps.

Seeds Aril absent. Testa pachychalazal, thick, vascularized. Tegmen absent. Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, with chlorophyll. Cotyledons two, thick, cordate, with backwardly directed lobes, with amyloid (xyloglucans). Germination phanerocotylar. Radicula ephemeral.

Phytochemistry Flavonols (kaempferol, quercetin, myricetin), cyanidin, delphinidin, ellagic acid, non-hydrolyzable tannins, caffeic acid and other polyphenols, glucosinolates (m-methoxybenzyl isothiocyanate) synthesized from phenylalanine and/or tyrosine, and erucic acid and other similar fatty acids (in seeds) present. Saponins and cyanogenic compounds not found. Carbohydrate stored as oligosaccharides with isokestose bonds.

Moringales R. Br. in C. F. P. von Martius, Consp. Regn. Veg.: 51. Sep-Oct 1835 [‘Moringeae’]

Distribution Semiarid regions in northeastern Africa, southern Angola, Namibia, southern Madagascar, southern Arabian Peninsula, southern Iran, Pakistan, India.

Habit Bisexual, deciduous trees or shrubs (sometimes pachycaul, sometimes stem succulents). Roots thick and fleshy, sometimes with root tubers.

Vegetative anatomy Phellogen? Primary medullary strands narrow. Medulla often with central lysigenous mucilage canals. Vessel elements with simple perforation plates; lateral pits alternate, simple or bordered pits. Vestured or non-vestured pits present. Imperforate tracheary xylem elements libriform fibres with simple pits, non-septate. Wood rays uniseriate to multiseriate, homocellular. Axial parenchyma paratracheal aliform, lozenge-aliform, confluent, or vasicentric. Wood elements partially storied. Sieve tube plastids S type. Nodes ?, multilacunar. Stem, leaves and flowers with myrosine cells and cells containing protein-rich cisternae from endoplasmic reticulum. Wood rays, phloem parenchyma and cortex with schizogenous intercellular ducts containing gums (rubber). Prismatic calciumoxalate crystals abundant. Cortex or xylem sometimes with druses. Phloem rays sometimes with crystals or sclereids.

Leaves Alternate (spiral), twice or three times imparipinnate, usually with persistent rhachis, with ? ptyxis. Stipules and stipulules as stalked glandular hairs or absent; leaf sheath absent. Petiole vascular bundle transection? Leaf base and articulations with distinct glands. Venation pinnate (to palmate). Stomata anomocytic. Cuticular wax crystalloids as rosettes of platelets (Fabales type). Epidermis often with mucilaginous idioblasts. Mesophyll with cells containing calciumoxalate druses. Leaflet margins entire. Extrafloral nectaries sometimes present between leaflets.

Flowers Usually transversely (obliquely) zygomorphic (sometimes actinomorphic). Hypanthium cupular or shortly tubular, often oblique (partially formed by inwardly bent receptacle), with basal whorl of nectaries. Hypogyny. Sepals five, with imbricate aestivation, petaloid, often unequal in size, free. Petals five, with imbricate aestivation; median petal adaxial and usually larger than recurved posterior petal pair, or abaxial; two lateral petals erect (anterior petal largest), free. Nectariferous disc annular, intrastaminal, at hypanthial base.

Androecium Stamens five, antepetalous-antesepalous. Filaments unequal in length, hairy at base, inserted at margin of nectariferous disc, free from each other and from tepals. Anthers declinate, dorsifixed, versatile?, usually disporangiate (rarely tetrasporangiate), introrse, longicidal (dehiscing by longitudinal slits; forming tube, through which style grows). Tapetum secretory. Staminodia three to five, extrastaminal, more or less antesepalous, alternating with fertile stamens.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3(–4)-colporate, shed as monads, usually bicellular (rarely tricellular) at dispersal. Exine tectate, with granular infratectum, sparsely perforate to punctate, psilate.

Gynoecium Pistil composed of (two or) three (or four) connate carpels. Ovary superior, unilocular, curved, stipitate (with gynophore). Style single, simple, narrow, curved, hollow (with stylar canal). Stigma capitate to truncate-porate (punctate?), hollow, type? Pistillodium absent.

Ovules Placentation parietal. Ovules numerous per carpel, anatropous, pendulous, apotropous?, bitegmic, crassinucellar. Micropyle bistomal to exostomal, Z-shaped (zig-zag). Outer integument ? cell layers thick, vascularized. Inner integument approx. three cell layers thick. Parietal tissue approx. three layers thick. Endothelium present. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis asterad.

Fruit Three- to twelve-angled narrowly elongate explosion (explosively dehiscent) loculicidal capsule.

Seeds Seeds usually three-angled to three-winged. Aril absent. Testa vascularized, multiplicative. Mesotesta thick, multiplicative; outer and inner parts of cell walls with spiral thickenings; middle part strongly thickened. Tegmen thin, usually not multiplicative. Exotegmen not fibrous. Perisperm not developed. Endosperm usually absent. Embryo large, straight, well differentiated, oily, with chlorophyll. Cotyledons two (or three). Hypocotyl swollen (often tuberous). Epicotyl sometimes swollen. Germination phanerocotylar or cryptocotylar. Leaves in young plants palmately compound or simple, palmately veined.

Phytochemistry Flavonols (kaempferol, quercetin) and glucosinolates (synthesized from phenylalanine and/or tyrosine, and from valine/isoleucine and/or leucine) and alkaloids present. Ellagic acid, tannins, proanthocyanidins, saponins, and cyanogenic compounds not found.

Use Ornamental plants (e.g. Moringa oleifera), vegetables, oil plants, medicinal plants, spices, water purification (seeds).

Distribution Tropical West and Central Africa, southwestern tropical Africa.

Vegetative anatomy Phellogen? Secondary lateral growth sometimes anomalous? Vessel elements with simple? perforation plates; lateral pits?, simple pits. Non-vestured pits present. Imperforate tracheary xylem elements ? with bordered pits. Wood rays? Axial parenchyma? Wood elements storied? Sieve tube plastids S type? Nodes 1:1, unilacunar with one leaf trace, or 3:3, trilacunar with three traces. Myrosin cells present. Crystals?

Leaves Alternate (spiral), simple, entire, with ? ptyxis. Stipules minute; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate. Stomata anomocytic. Cuticular wax crystalloids? Epidermis with mucilaginous idioblasts. Leaf margin entire.

Flowers Actinomorphic to bisymmetric. Hypogyny. Sepals five, with valvate aestivation, sometimes persistent, free or somewhat connate at base. Petals five, with imbricate aestivation, often clawed, arched below around nectaries/androgynophore and connivent through lanate hairs at edges. Nectariferous disc extrastaminal, annular, thick, fleshy, as short androgynophore.

Androecium Stamens (nine or) ten (to 13), diplostemonous or triplostemonous. Filaments filiform, branched or simple?, free or connate at base, free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits); connective slightly prolonged. Tapetum secretory? Female flowers with ten staminodia.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolporate, shed as monads, tricellular? at dispersal. Exine semitectate, with columellate infratectum, microreticulate.

Gynoecium Pistil composed of (three to) five connate antesepalous carpels. Ovary superior, (trilocular to) quinquelocular, shortly stipitate (gynophore). Style single, simple, long. Stigma short, (trilobate to) quinquelobate, type? Male flowers with pistillodium.

Ovules Placentation axile. Ovules approx. ten per carpel (in two or three rows), anatropous?, bitegmic, crassinucellar? Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A many-seeded berry (containing brazzein, a protein with extremely sweet taste).

Seeds Aril absent. Testa lanate, with external layer of white, wooly, elongate cells. Tegmen? Perisperm not developed. Endosperm absent. Embryo strongly curved, without chlorophyll. Cotyledons two. Germination?

Phytochemistry Insufficiently known. Alkaloids, benzyl and 4-methoxybenzyl glucosinolates synthesized from phenylalanine and/or tyrosine, saponins, sweet-tasting proteins (brazzein, c. 2.000 times sweeter than saccharose, pentadin) present. Ellagic acid? Tannins?

Resedales Bercht. et J. Presl, Přir. Rostlin: 218. Jan-Apr 1820 [‘Resedaceae’]; Astrocarpaceae A. Kern., Pflanzenleben 2: 688. 6-13 Jun 1891 [’Asterocarpaceae’]; Resedineae Engl., Syllabus, ed. 2: 123. Mai 1898; Stixaceae Doweld, New Syllabus Pl. Fam.: 746. Apr 2007

Distribution Mostly dry areas in the southwestern United States, Mexico, Central America, the West Indies, Macaronesia, northern and eastern Africa, South Africa, the Mediterranean, Europe (except northern parts), the Arabic Peninsula, Socotra, western Asia to northwestern India, western Central Asia, western Siberia, southeastern Himalayas, Hainan, Southeast Asia, Malesia.

Habit Usually bisexusal (in ‘Ochradenus’ sometimes androdioecious, in Forchhammeria dioecious), evergreen or deciduous shrubs or suffrutices, or perennial, biennial or annual herbs, rarely climbing. Some species are xerophytes.

Vegetative anatomy Roots rarely with mycorrhiza. Phellogen? Secondary lateral growth sometimes anomalous from successive cambia. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured and non-vestured pits present. Imperforate tracheary xylem elements libriform fibres with usually simple (sometimes bordered) pits, non-septate. Wood rays uniseriate or multiseriate, homocellular or heterocellular. Axial parenchyma apotracheal diffuse, or paratracheal scanty, confluent, vasicentric, or marginal-banded. Wood elements not storied. Intraxylary phloem sometimes concentric. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace. Myrosin cells idioblastic. Stem and foliar epidermis sometimes with mucilaginous idioblasts. Crystals? Endoplasmic reticulum with ER-dependent cisternae.

Trichomes Hairs usually unicellular (in Tirania multicellular, uniseriate), simple; glandular hairs absent?

Leaves Alternate (spiral), simple or pinnately compound (rarely trifoliolate), entire or pinnately lobed, with ? ptyxis. Stipules small, intrapetiolar, often as glands, or absent; leaf sheath absent (in Tirania as spines). Petiole vascular bundle transection? Venation pinnate (rarely palmate) or leaves one-veined. Stomata anomocytic. Cuticular wax crystalloids? Stomatal myrosin cells absent. Epidermis with or without mucilaginous idioblasts. Mesophyll usually without calciumoxalate crystals (in Forchhammeria with sclerenchymatous cells). Leaf margin sinuate or entire. In Caylusea large water absorbing hairs present along leaf margin.

Inflorescence Usually terminal (sometimes axillary), simple or compound raceme, spike or panicle (in Forchhammeria trifoliata sometimes cymose; flowers in Tirania solitary axillary). Floral prophylls (bracteoles) absent.

Flowers Zygomorphic, usually small. Hypogyny or half epigyny. Sepals (four to) six (to eight), with valvate or somewhat imbricate aestivation, persistent, usually free (occasionally somewhat connate at base); sepals in Tirania six, in Forchhammeria numerous (bracts?) and in Stixis three to numerous, with imbricate aestivation. Petals (two or four to) six (to eight), with valvate or open aestivation, unequally sized (two adaxial petals largest, lateral petals smaller and less split, abaxial petals smallest), clawed, caducous or persistent, usually free (petals absent in ‘Ochradenus’), usually with adaxial ligule (absent in ‘Oligomeris’) inserted between claw and apically usually fringed limb; petals in Tirania six, with imbricate aestivation, absent in Forchhammeria and Stixis. Gynophore or androgynophore present in some species; androgynophore present in Stixis. Nectariferous disc extrastaminal (or sometimes absent), often widest adaxially, with nectariferous gland on abaxial side; nectary sometimes petaloid (nectariferous disc absent in ‘Oligomeris’).

Androecium Stamens (three to) 16 to 22 (to more than 50; in ’Ochradenus’ numerous; in ’Oligomeris’ linifolia three), usually in one whorl (sometimes in several whorls), centrifugally developing from annular primordium. Filaments usually free (sometimes connate at base), free from tepals; filaments in Forchhammeria inserted on annular outgrowth surrounding base of androgynophore, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits); anthers in Forchhammeria with short unicellular hairs provided with cuticular longitudinal folds, dorsifixed to basifixed, often versatile, tetrasporangiate, latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenes simultaneous. Pollen grains tricolpate, tricolporate or tricolporoidate, shed as monads, bicellular at dispersal; pollen grains in Stixis (2–)3(–4)-colporoidate and very small, in Forchhammeria tricolporate. Exine tectate or semitectate, with columellate infratectum, reticulate, microreticulate or microperforate, foveolate, striate or rugulate.

Gynoecium Pistil composed of (two or) three to six (to eight), connate or apically and adaxially often incompletely paracarpously connate, antesepalous carpels (in ’Ochradenus’ finally closed); when carpels three, then median carpel often adaxial (carpels in Caylusea connate only at base; carpels in Sesamoides free, apocarpy). Ovary superior or semi-inferior, usually unilocular (sometimes bilocular to multilocular), shortly stipitate (gynophore and sometimes androgynophore); one locule and one seed in Forchhammeria usually degenerating; ovary sessile in Tirania. Style usually absent (sometimes single, simple or branched). Stigmas two to five, usually small and narrow (in Forchhammeria fleshy), commissural and/or dorsal, non-papillate, Dry type. Pistillodium absent.

Ovules Placentation usually parietal (in e.g. Tirania axile; in Caylusea basal; in Sesamoides laminal). Ovules two to numerous (in Forchhammeria and Sesamoides usually one, rarely two) per carpel, anatropous (to hemianatropous?) to campylotropous, pendulous or ascending, bitegmic, usually crassinucellar (rarely tenuinucellar). Micropyle usually endostomal (sometimes bistomal). Outer integument approx. two cell layers thick. Inner integument three or four cell layers thick. Hypostase present. Archespore multicellular. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis onagrad.

Fruit Usually a soft, thin-walled one- to many-seeded capsule apically open between stigmas (in Sesamoides an assemblage of follicles, carpidia; in some species of ’Ochradenus’ baccate with fleshy pericarp).

Seeds Aril usually present, formed from apex of outer integument. Elaiosome (carunculi?; lipid-rich tissue adjacent to hilum) present in some species. Testa non-multiplicative. Exotesta? Endotestal cells cuboid, with thickened unlignified walls and often crystalliferous. Tegmen non-multiplicative. Exotegmic cells fibrous, with lignified walls. Endotegmen crystalliferous. Perisperm not developed. Endosperm sparse or absent. Embryo large, curved or plicate, well differentiated, oily, with chlorophyll (at least in some species of Reseda). Cotyledons two, sometimes unequal (to very unequal) in size. Germination phanerocotylar, hypogeous or epigeous.

Phytochemistry Flavonols (kaempferol, quercetin), methylated flavonols, cyanogenic compounds, glucosinolates (synthesized from phenylalanine and/or tyrosine), and aromatic m-carboxycinnamic acids present. Ellagic acid, tannins, and proanthocyanidins not found. BCAA glucosinolates absent. Sinapic acid not accumulated. Methyl glucosinolates (synthesized from phenylalanine and/or tyrosine and methionin?) present in Forchhammeria.

Use Ornamental plants, vegetables (Caylusea abyssinica), dyeing sources (Reseda luteola), perfumes (Reseda odorata).

Azimaceae Wight et Gardner in Calcutta J. Nat. Hist. 6: 52. Apr 1845; Salvadorineae Engl., Syllabus, ed. 2: 171. Mai 1898; Salvadorales R. Dahlgren ex Reveal in Phytologia 74: 174. 25 Mar 1993

Distribution Africa, Madagascar, the southern Arabian Peninsula, Pakistan, India, Sri Lanka, southern China, Southeast Asia, West Malesia, the Philippines, often in drier regions.

Habit Bisexual, polygamomonoecious, dioecious, or polygamodioecious, evergreen shrubs (sometimes climbing) or small trees. Xerophytes, sometimes spiny.

Vegetative anatomy Phellogen ab initio superficial. Primary vascular tissue a cylinder of vascular bundles. Secondary lateral growth anomalous, from simple cambial cylinder. Vessel elements with simple perforation plates; lateral pits alternate, bordered pits. Vestured pits present in Salvadora. Imperforate tracheary xylem elements libriform fibres with simple pits, non-septate. Wood rays uniseriate or multiseriate, homocellular (or slightly heterocellular). Axial parenchyma paratracheal scanty vasicentric, or banded. Wood elements partially storied. Intraxylary (concentric or diffuse) phloem present in Dobera and Salvadora. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace (Salvadora), or 1:2, unilacunar with two traces (Azima, Salvadora). Myrosin cells absent. Prismatic calciumoxalate crystals abundant. Wood rays with rhomboidal calciumoxalate crystals. Endoplasmic reticulum with proteinaceous vacuole-like dilated cisternae.

Leaves Opposite, simple, entire, usually coriaceous, with flat to curved ptyxis (Salvadora). Stipules very small; leaf sheath absent. Petiole vascular bundle transection? Venation pinnate. Stomata usually paracytic (sometimes anomocytic or anisocytic). Cuticular wax crystalloids as platelets. Mesophyll with cells containing calciumoxalate druses. Leaf margin entire.

Inflorescence Terminal or axillary, simple raceme or compound, fasciculate inflorescence. Floral prophylls (bracteoles) usually present.

Flowers Usually actinomorphic (occasionally somewhat zygomorphic), obliquely orientated, small. Short hypanthium present in Salvadora. Hypogyny. Sepals two to four (or five), with imbricate to contorted aestivation, connate at base into tube. Petals two to four (or five), with imbricate to contorted aestivation, often with glands (staminodia?) on adaxial side, free (Azima, Dobera) or connate at base (Salvadora). Antepetalous nectariferous glands (probably not staminal) alternating with stamens (in Dobera extrastaminal, in Salvadora interstaminal) or absent (Azima).

Androecium Stamens two to four (or five), haplostemonous, antesepalous, alternipetalous. Filaments free (Azima) or connate at base (Dobera), free from or adnate at base to petals (Salvadora). Anthers dorsifixed, versatile?, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent (or possibly as glands?).

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporoidate (sometimes hexacolporoidate), shed as monads, bicellular at dispersal. Exine tectate, with columellate infratectum, punctate, perforate, reticulate or striate.

Gynoecium Pistil composed of two connate carpels, in Salvadora obliquely orientated. Ovary superior, bilocular (Azima) or unilocular (pseudomonomerous; Dobera, Salvadora); in Dobera and Azima apically quadrilocular due to apical secondary septum; ovary shortly stipitate (gynophore) in Salvadora. Style single, simple, short. Stigma capitate (Salvadora) or slightly bilobate (Azima, Dobera), type? Male flowers with pistillodium.

Ovules Placentation basal-parietal (when ovary unilocular) or axile-basal (when ovary bilocular). Ovules one or two per carpel, anatropous, ascending, apotropous, bitegmic, crassinucellar. Micropyle usually endostomal (sometimes exostomal). Outer integument approx. ten to 15 cell layers thick (in Azima vascularized). Inner integument three to five cell layers thick. Obturator dome-shaped, chalazal, present in Salvadora (absent in Azima and Dobera). Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Endosperm haustoria absent. Embryogenesis asterad.

Seeds Aril absent. Seed coat exotestal. Testa multiplicative. Exotestal cells palisade, somewhat thickened, crystalliferous, with inner walls mucilaginous. Endotesta present. Tegmen non-multiplicative. Exotegmen fibrous or non-fibrous, finally crushed; exotegmic cell walls unlignified. Endotegmen crushed. Perisperm not developed. Endosperm absent. Embryo straight, well differentiated, chlorophyll? Cotyledons two, cordate, thick, plano-convex, oily. Germination?

Phytochemistry Flavonols (quercetin), piperidine alkaloids, mustard oils based on glucotropaeolin, and cyanogenic compounds present. Ellagic acid, tannins, and proanthocyanidins not found.

Habit Bisexual, evergreen shrub with pungent smell and well differentiated long and short shoots. Xerophyte.

Vegetative anatomy Phellogen ab initio superficial. Primary medullary rays narrow. Primary vascular tissue cylinder, without separate vascular bundles. Vessel elements with simple perforation plates; lateral pits alternate. Non-vestured pits present. Imperforate tracheary xylem elements fibre tracheids with bordered pits, non-septate? (also vasicentric tracheids). Wood rays uniseriate. Axial parenchyma usually paratracheal scanty (sometimes apotracheal diffuse or diffuse-in-aggregates, or banded). Wood elements not storied. Sieve tube plastids S type? Nodes 1:1, unilacunar with one leaf trace. Myrosin cells absent. Crystals rare (rhomboidal) or absent; druses present in cortical parenchyma cells.

Trichomes Unicellular T-shaped malpighiacean hairs, situated on basal multicellular podium, with branches parallel to longitudinal axis of organ (as in Brassicaceae).

Leaves Alternate (spiral), simple, entire, isobifacial, often coriaceous, with ? ptyxis. Stipules and leaf sheath absent. Petiole vascular bundle transection? Venation pinnate or palmate; secondary veins subbasal (inserted near leaf base). Stomata anomocytic. Cuticular wax crystalloids? Leaf margin entire.

Flowers Actinomorphic, large. Receptacle elongated into short androphore and relatively short gynophore. Hypogyny. Sepals (five or) six (or seven), with valvate? aestivation, swollen at base, connate into calyptra and at anthesis irregularly dehiscing with one or two valves, caducous or persistent. Petals (five or) six (or seven), with imbricate aestivation, clawed, caducous, free. Nectary absent? Disc absent.

Androecium Stamens (c. 40 to) c. 60 to 76 (from six staminal primordia), in five to seven vertical alternipetalous rows of staminal pairs on receptacle, centrifugally developing. Filaments free from each other (seemingly?), free from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory? Staminodia absent.

Pollen grains Microsporogenesis simultaneous? Pollen grains tricolpate, shed as monads, ?-cellular at dispersal. Exine semitectate, with columellate infratectum, striato-rugulate.

Gynoecium Pistil composed of three carpels, connate along their prolonged adaxial margins; commissural fusion of marginal ventral carpellary vascular bundles. Ovary superior, trilocular. Style single, simple, short, trifid. Stigmas subcapitate, type? Pistillodium absent.

Ovules Placentation axile. Ovules ten to 14 per carpel, arranged in two rows, anatropous, bitegmic, crassinucellar. Micropyle ?-stomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development? Endosperm haustoria? Embryogenesis?

Fruit A septifragal capsule, dehiscing with three valves and with persistent thin central columella corresponding to replum.

Seeds Seed at cotyledonary end aleate, stellate in transverse section. Aril absent. Testa multiplicative?, soft, thin, aleately expanded at chalazal end. Tegmen multiplicative? Exotegmen non-fibrous. Endotegmen? Perisperm not developed. Endosperm sparse, one cell layer thick. Embryo flat, straight, chlorophyll? Cotyledons two, cordate. Germination phanerocotylar.

Phytochemistry Uniknown. Glucosinolates synthesized from phenylalanine and/or tyrosine?

Tovariales Nakai, Chosakuronbun Mokuroku [Ord. Fam. Trib. Nov.]: 239. 20 Jul 1943

Habit Bisexual, annual or perennial herbs, or evergreen shrubs or suffrutices (sometimes climbing or arborescent). Leaves when dry with coumarin-like scent.

Vegetative anatomy Phellogen? Vessel elements with simple perforation plates; lateral pits alternate or scalariform, simple pits. Non-vestured pits present. Imperforate tracheary xylem elements libriform fibres with simple pits, septate. Wood rays uniseriate or multiseriate, heterocellular? Axial parenchyma scanty vasicentric. Wood elements not storied. Sieve tube plastids S type. Nodes 1:1, unilacunar with one leaf trace? Myrosin cells idioblastic or stomatal. Crystals? Cells with organelle-like expanded ER-cisternae absent.

Leaves Alternate (spiral), trifoliolate, with ? ptyxis. Stipules cauline or present on leaf base, minute, early caducous; leaf sheath absent. Petiole vascular bundle transection arcuate. Venation palmate. Stomata anomocytic. Cuticular wax crystalloids? Leaflet margins entire.

Flowers Bisymmetric, small. Hypogyny. Sepals (six to) eight (to ten), with imbricate or open aestivation, in one whorl, caducous, free. Petals (six to) eight (or nine), with imbricate aestivation, in one whorl, often shortly clawed, free. Nectariferous disc largely extrastaminal, lobate, with nectaries between staminal bases.

Androecium Stamens (six to) eight (to ten), haplostemonous, antesepalous, alternipetalous. Filaments widened at base, sometimes hairy, free from each other and from tepals. Anthers with short unicellular hairs provided with cuticular longitudinal folds, basifixed, non-versatile, tetrasporangiate, introrse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent.

Pollen grains Microsporogenesis simultaneous. Pollen grains (2–)3-colpor(oid)ate, shed as monads, bicellular at dispersal. Exine semitectate, with columellate infratectum, reticulate.

Gynoecium Pistil composed of (five or) six (to eight) connate alternisepalous carpels. Ovary superior, incompletely sexalocular (sometimes quinque-, septa- or octalocular), sometimes stipitate (gynophore short or absent). Style single, simple, short, persistent, or absent. Stigma peltate or sexalobate to octalobate, type? Pistillodium absent.

Ovules Placentation axile in lower part of ovary to intrusively parietal in upper part. Ovules numerous per carpel, in several rows, ab initio anatropous, finally (after fertilization) campylotropous, bitegmic, crassinucellar. Funicle long. Micropyle bistomal, Z-shaped (zig-zag). Outer integument approx. two cell layers thick. Inner integument approx. three cell layers thick. Megagametophyte monosporous, Polygonum type. Synergids with a filiform apparatus. Endosperm development ab initio nuclear. Endosperm haustoria? Embryogenesis?

Seeds Aril absent. Seed coat testal-tegmic? Testa non-multiplicative. Exotestal cells enlarged, with tannins and thickened walls. Endotestal cells small. Tegmen non-multiplicative. Exotegmic cells fibrous, with reticulately thickened walls. Endotegmen tanniniferous. Perisperm not developed. Endosperm sparse, thin, oily. Embryo curved along periphery of seed, well differentiated, oily, chlorophyll? Cotyledons two, flattened. Germination?

Phytochemistry Insufficiently known. Glucosinolates (not synthesized from phenylalanine or tyrosine) and cyanogenic compounds present. Tannins?

Tropaeolales Bercht. et J. Presl, Přir. Rostlin: 220. Jan-Apr 1820 [‘Tropaeoleae’]

Distribution Mountain areas from southern Mexico to Tierra del Fuego, southern Brazil to southern Argentina.

Habit Bisexual, perennial or annual herbs, usually climbing with petioles or pedicel tendrils, more or less fleshy (sometimes slightly succulent). Several species form root or stem tubers from rhizome or from hypocotyl.

Vegetative anatomy Phellogen ab initio superficial to deeply seated. Primary vascular tissue cylinder of separate vascular bundles. Endodermis sometimes significant. Pericyclic fibres absent. Vessels present in root and stem. Vessel elements with usually simple (sometimes reticulate, scalariform or intermediate between scalariform and multiperforate) perforation plates; lateral pits alternate, simple pits. Non-vestured pits present. Imperforate tracheary xylem elements usually tracheids (libriform fibres present in stem; also scarce vasicentric tracheids) with simple or bordered pits, non-septate. Wood rays uniseriate (in root) or multiseriate, heterocellular. Axial parenchyma paratracheal scanty vasicentric. Wood elements not storied. Sieve tube plastids Ss type. Nodes 3:3, trilacunar with three leaf traces. Myrosine cells only idioblastic. Crystals?

Trichomes Hairs multicellular (few-celled), uniseriate, simple, or absent; glandular hairs sometimes present.

Leaves Usually alternate (sometimes opposite in lower parts), usually simple (sometimes palmately compound), entire or palmately lobed, often peltate, with flat ptyxis. Stipules small, fimbriate or more or less foliaceous or absent (often present only in seedlings), usually caducous; leaf sheath absent. Petiole often climbing. Petiole vascular bundle transection annular. Venation palmate. Stomata anomocytic, without subsidiary cells. Cuticular wax crystalloids as clustered tubuli (Berberis type), chemically dominated by nonacosan-10-ol. Lamina sometimes gland-dotted. Epidermis sometimes with mucilaginous idioblasts. Myrosine cells containing myrosinase abundant; stomatal myrosin cells absent. Leaf margin serrate or entire.

Inflorescence Flowers usually axillary solitary (rarely in axillary umbel-like inflorescence). Floral prophylls (bracteoles) present or absent.

Flowers Zygomorphic, often large. Hypogyny. Sepals five, with imbricate or valvate aestivation, petaloid; adaxial, or adaxial and two lateral sepals expanded into nectariferous spur (rarely absent), usually free (sometimes three connate), caducous. Petals usually two upper adaxial and three lower abaxial (lower petals often absent), with imbricate aestivation, often clawed, free. Nectary extrastaminal, continuing into spur. Disc absent. Hairs on lower margin of abaxial (anterior) petals forming barrier against small non-pollinating insects (nectar thieves).

Androecium Stamens 4+4 (two antepetalous – median – stamens suppressed). Filaments filiform, free from each other and from tepals. Anthers basifixed, non-versatile, tetrasporangiate, introrse-latrorse, longicidal (dehiscing by longitudinal slits). Tapetum secretory. Staminodia absent. During anthesis in Tropaeolum majus, stamens bend forth successively one after the other to nectariferous spur liberating their pollen grains and subsequently returning to their original position; style finally bending against spur.

Pollen grains Microsporogenesis simultaneous. Pollen grains usually tricolporate (sometimes dicolporate), shed as monads, tricellular at dispersal. Exine semitectate, with columellate infratectum, reticulate (sometimes partially striate).

Gynoecium Pistil composed of three connate carpels; median carpel adaxial. Ovary superior, trilocular. Style single, narrow, short. Stigma trilobate, non-papillate, Dry type. Pistillodium absent.

Ovules Placentation apical-axile. Ovules two per carpel (one degenerating), anatropous, pendulous, epitropous, bitegmic, tenuinucellar, pachychalazal (massive chalaza becoming meristematic and forming seed coat). Micropyle endostomal. Outer integument ? cell layers thick. Inner integument ? cell layers thick. Megagametophyte monosporous, Polygonum type. Endosperm development ab initio nuclear. Two uppermost cells of proembryo several times dividing and forming basal system of haustoria (at least in Tropaeolum majus). Peripheral cells of basal cell mass (adjacent to suspensor and on opposite side of funicle) forming rhizomorphic suspensor haustorium, penetrating integument below micropyle, growing around ovule and into carpellary wall. Additional suspensor haustorium developing from peripheral cells of basal cell mass adjacent to funicle, growing through integument and funicle and reaching placental vascular bundle, growing further along placenta. Embryogenesis solanad.

Fruit A schizocarp with usually three (rarely one) drupaceous or nutlike mericarps (rarely a samara). Carpophore rarely present.

Seeds Seeds pachychalazal (seed coat formed by chalaza). Aril absent. Testa multiplicative, indistinct (outer integument disappearing during maturation). Part of mesotesta suberized. Endotesta (inner epidermis) lacking crystalliferous and sclerotic cells. Tegmen non-multiplicative. Exotegmen not developing into sclerotic layer. Perisperm not developed. Endosperm absent. Embryo large, straight, well differentiated, with chlorophyll. Cotyledons two, thick, plano-convex, with amyloid (xyloglucans). Germination cryptocotylar.

Phytochemistry Flavonols (kaempferol, quercetin), benzyl and 4-methoxybenzyl glucosinolates (synthesized from phenylalanine and/or tyrosine and valine/isoleucine and/or leucine), erucic acid, eicosenoic acid, and cucurbitacins present. Ellagic acid, proanthocyanidins and cyanogenic compounds not found.

Use Ornamental plants, vegetables (Tropaeolum tuberosum etc.), pickled unripe fruits.

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Al-Shehbaz IA. 1990d. A note on the Chilean endemic Draba thlaspiformis (Brassicaceae). – J. Arnold Arbor. 71: 385-387.

Al-Shehbaz IA. 1991. Novelties in Draba (Brassicaceae) from Venezuela, Ecuador, and Peru. – Novon 1: 67-70.

Al-Shehbaz IA. 1992. Draba barclayana (Brassicaceae), a new species from Colombia. – Novon 2: 4-5.

Al-Shehbaz IA. 1994a. Petroravenia (Brassicaceae), a new genus from Argentina. – Novon 4: 191-196.

Al-Shehbaz IA. 1994b. Three new South American species of Draba (Brassicaceae). – Novon 4: 197-202.

Al-Shehbaz IA. 1999. Generic placement of species excluded from Arabidopsis (Brassicaceae). – Novon 9: 296-307.

Al-Shehbaz IA. 2000a. A revision of Pegaeophyton (Brassicaceae). – Edinburgh J. Bot. 57: 157-170.

Al-Shehbaz IA. 2000b. Baimashania (Brassicaceae), a new genus from China. – Novon 10: 320-322.

Al-Shehbaz IA. 2000c. What is Nasturtium tibeticum (Brassicaceae)? – Novon 10: 334-336.

Al-Shehbaz IA. 2001. A review of gamosepaly in the Brassicaceae and a revision of Desideria, with a critical evaluation of related genera. – Ann. Missouri Bot. Gard. 87: 549-563.

Al-Shehbaz IA. 2002a. Noccaea nepalensis, a new species from Nepal, and four new combinations in Noccaea (Brassicaceae). – Adansonia, sér. III, 24: 89-92.

Al-Shehbaz IA. 2002b. New species of Alyssum, Aphragmus, Arabis, and Sinosophiopsis (Brassicaceae) from China and India. – Novon 12: 309-313.

Al-Shehbaz IA. 2002c. Six new species of Draba (Brassicaceae) from the Himalayas. – Novon 12: 314-318.

Al-Shehbaz IA. 2003a. Proposal to conserve the name Smelowskia against Redowskia (Brassicaceae). – Taxon 52: 360-361.

Al-Shehbaz IA. 2003b. Transfer of most North American species of Arabis to Boechera (Brassicaceae). – Novon 13: 381-391.

Al-Shehbaz IA. 2003c. A synopsis of Tropidocarpum (Brassicaceae). – Novon 13: 392-395.

Al-Shehbaz IA. 2003d. Aphragmus bouffordii, a new species from Tibet and a synopsis of Aphragmus (Brassicaceae). – Harvard Pap. Bot. 8: 25-27.

Al-Shehbaz IA. 2004a. Novelties and notes on miscellaneous Asian Brassicaceae. – Novon 14: 153-157.

Al-Shehbaz IA. 2004b. Two new species of Draba (Brassicaceae): D. miehorum from Tibet and D. sagasteguii from Peru. – Novon 14: 249-252.

Al-Shehbaz IA. 2004c. A synopsis of the South American Neuontobotrys (Brassicaceae). – Novon 14: 253-257.

Al-Shehbaz IA. 2004d. A synopsis of the South American Weberbauera (Brassicaceae). – Novon 14: 258-268.

Al-Shehbaz IA. 2005a. Proposal to conserve the name Erucastrum against Kibera and Hirschfeldia (Brassicaceae). – Taxon 54: 204-205.

Al-Shehbaz IA. 2005b. Nomenclatural notes on Eurasian Arabis (Brassicaceae). – Novon 15: 519-524.

Al-Shehbaz IA. 2005c. Hesperidanthus (Brassicaceae) revisited. – Harvard Pap. Bot. 10: 47-51.

Al-Shehbaz IA. 2006a. The genus Sisymbrium in South America, with synopses of the genera Chilocardamum, Mostacillastrum, Neuontobotrys, and Polypsecadium (Brassicaceae). – Darwiniana 44: 341-358.

Al-Shehbaz IA. 2006b. New or noteworthy taxa of Argentinean and Chilean Brassicaceae (Cruciferae). – Darwiniana 44: 359-362.

Al-Shehbaz IA. 2007a. Two new species of Draba and Eutrema (Brassicaceae) from Sichuan, China. – Harvard Pap. Bot. 11: 277-279.

Al-Shehbaz IA. 2007b. Generic limits of Dryopetalon, Rollinsia, Sibara, and Thelypodiopsis (Brassicaceae), and a synopsis of Dryopetalon. – Novon 17: 397-402.

Al-Shehbaz IA. 2007c. Arabis elgonensis (Brassicaceae), a new species from Mount Elgon, Uganda. – Harvard Pap. Bot. 12: 387-388.

Al-Shehbaz IA. 2007d. The North American genus Sandbergia (Boechereae, Brassicaceae). – Harvard Pap. Bot. 12: 425-427.

Al-Shehbaz IA. 2010a. The legacy of Asa Gray’s South American botany: examples of amphitropical disjunctions from the Brassicaceae. – Harvard Pap. Bot. 15: 205-208.

Al-Shehbaz IA. 2010b. Ivania juncalensis, a second species in the Chilean endemic Ivania (Brassicaceae). – Harvard Pap. Bot. 15: 343-345.

Al-Shehbaz IA. 2010c. A synopsis of the genus Sibara (Brassicaceae). – Harvard Pap. Bot. 15: 139-147.

Al-Shehbaz IA. 2010d. Menonvillea zuloagaensis and Mostacillastrum hunzikeri (Brassicaceae), two new species from Argentina. – Darwiniana 48: 59-63.

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Al-Shehbaz IA. 2012b. Notes on miscellaneous species of the tribe Thelypodieae (Brassicaceae). – Harvard Pap. Bot. 17: 3-10.

Al-Shehbaz IA. 2013. Clypeola is united with Alyssum (Brassicaceae). – Harvard Pap. Bot. 18: 125-128.

Al-Shehbaz IA. 2016. A revision of the Mexican endemic Asta (Brassicaceae). – Novon 25: 8-11.

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Al-Shebaz IA, German DA. 2016. Three new genera in the tribe Euclidieae (Brassicaceae). – Novon 25: 12-17.

Al-Shehbaz IA, Guang Y. 2000a. A reconsideration of the genus Eurycarpus (Brassicaceae). – Novon 10: 346-348.

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Al-Shehbaz IA, Iltis H. 1993. Romanschulzia mexicana (Brassicaceae), a remarkable new species from Guerrero Mexico. – Novon 3: 96-98.

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Al-Shehbaz IA, Mummenhoff K. 2011. Stubendorffia and Winklera belong to the expanded Lepidium (Brassicaceae). – Edinburgh J. Bot. 68: 165-171.

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Al-Shehbaz IA, An ZY, Yang G. 1999. A revision of Sisymbriopsis (Brassicaceae). – Novon 9: 308-312.

Al-Shehbaz IA, Arai K, Ohba H. 1999. A new species of Hemilophia (Brassicaceae) from China. – Novon 9: 8-10.

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CAPPARALES Juss. ex Bercht. et J. Presl

AKANIACEAE Stapf

BATACEAE Mart. ex Perleb

BORTHWICKIACEAE Su, Wang, Zhang et Chen

BRASSICACEAE Burnett

CARICACEAE Dumort.

EMBLINGIACEAE (Pax) Airy Shaw

GYROSTEMONACEAE (Endl.) A. Juss.

KOEBERLINIACEAE Engl.

LIMNANTHACEAE R. Br.

MORINGACEAE Martinov

PENTADIPLANDRACEAE Hutch. et Dalziel

RESEDACEAE Martinov

SALVADORACEAE Lindl.

SETCHELLANTHACEAE Iltis

TOVARIACEAE Pax

TROPAEOLACEAE Juss. ex DC.