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FOC | Family List | FOC Vol. 8

Brassicaceae Burnett

十字花科 shi zi hua ke

Authors: Tai-yien Cheo, Lianli Lu, Guang Yang, Ihsan Al-Shehbaz & Vladimir Dorofeev

Lepidium chalepense

Credit: Photo: Al-Shehbaz

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

About 330 genera and 3500 species: all continents except Antarctica, mainly in temperate areas, with highest diversity in Irano-Turanian, Mediterranean, and W North American regions; 102 genera (eight endemic) and 412 species (115 endemic) in China.

Because of lack of a comprehensive classification based on phylogenetic relationships among genera, and in order to facilitate direct comparison between the accounts in FOC and FRPS, the sequence of genera follows Schulz (Nat. Pflanzenfam. 17b: 227-658. 1936). However, it should be emphasized that his system is largely artificial because he placed closely related genera in different tribes and remotely related genera in the same tribe. For example, Arabis, Aubrieta, Draba, and Macropodium, which are very closely related on the basis of recent molecular studies, were placed by Schulz in four different tribes, whereas the unrelated Capsella, Lepidium, and Thlaspi were placed in one tribe. The delimitation of genera is often difficult, and mature fruits are needed for reliable identification.

The Brassicaceae include many important crop plants that are grown as vegetables (Brassica, Nasturtium, Raphanus) and sources of vegetable oils (Brassica) and condiments (Armoracia, Brassica, Eutrema, Sinapis). Oils of Brassica probably rank first in terms of tonnage of the world's production of edible oils. The family includes many ornamentals in the genera Erysimum, Iberis Linnaeus, Lobularia, Malcolmia, and Matthiola. Of these, only Lobularia maritima has become naturalized in China. The family also includes more than 120 species of weeds. Arabidopsis thaliana, which is naturalized in China, has become the model organism in many fields of experimental biology.

The delimitation of genera in the Brassicaceae is often difficult because of the frequent independent evolution of what appear to be similar character states, the variability of a given character in one genus and its fixture in another, and the inadequate sampling of material by most authors. Fruit characters are essential in the identification of genera. However, a key emphasizing flowering material is given in addition to the one emphasizing fruit. The most reliable determination of genera can be achieved when the material has both fruit and flowers and when both keys are successfully used to reach the same genus.

The types of cotyledonary position, which in reality is the position of the radicle in relation to the cotyledons (e.g., radicle accumbent, incumbent, or conduplicate, though radicle referred to as "cotyledon" throughout) are important in the separation of many genera, and a cross section of the seed provides the easiest and simplest way of determining that position. The exact number of ovules per ovary can easily be determined from the fruit because aborted ovules persist through fruit dehiscence. However, it is more laborious to determine the ovule number from pistils and young fruit.

Cheo Tai-yien, Guo Rong-lin, Lan Young-zhen, Lou Lian-li, Kuan Ke-chien & An Zheng-xi. 1987. Cruciferae. In: Cheo Tai-yien, ed., Fl. Reipubl. Popularis Sin. 33: 1-483.

Leads marked with a "double asterisk" (**) indicate that mature fruits are necessary for the identification of genera in the subordinate couplet(s).


Key to fruiting material

1 Fruit silicles (less than 3 × longer than broad).   (2)
+ Fruit siliques (more than 3 × longer than broad).   (59)
       
2 (1) Fruit much wider than long, 2-16-spined, with prominently 3-20-veined lateral wings larger than the seed-bearing part   15 Pugionium
+ Fruit longer than wide or about as long as wide, not spiny, wingless or with veinless wings smaller than the seed-bearing part.   (3)
       
3 (2) Fruit flattened at a right angle to septum (angustiseptate).   (4)
+ Fruit flattened parallel to septum (latiseptate), terete, or angled.   (21)
       
4 (3) At least some of the trichomes branched.   (5)
+ Trichomes absent or exclusively simple.   (8)
       
5 (4) Cauline leaves sessile, auriculate or sagittate at base; fruit obdeltoid to obdeltoid-obcordate   19 Capsella
+ Cauline leaves petiolate or cuneate to petiolelike base, not auriculate or sagittate; fruit oblong, elliptic, obovate, or suborbicular.   (6)
       
6 (5) Fruiting pedicels strongly recurved, often forming a loop; fruit 1- or 2-seeded, often aborted on 1 side; plants rhizomatous perennials   23 Dipoma
+ Fruiting pedicels straight, erect, ascending, or divaricate; fruit more than 5-seeded, not aborted on 1 side; plants annual, biennial, or rarely non-rhizomatous perennials.   (7)
       
7 (6) Fruiting raceme bracteate throughout or rarely along proximal half; leaves 1- or 2-pinnatisect; fruit more than 20-seeded   20 Hedinia
+ Fruiting racemes ebracteate; leaves entire, dentate, or pinnatifid; fruit less than 20-seeded   21 Hornungia
       
8 (4) Seeds (or ovules) 4-24 per fruit.   (9)
+ Seeds (or ovules) 1 or 2 per fruit.   (13)
       
9 (8) Fruit borne on solitary pedicels originating from basal rosette   44 Pegaeophyton
+ Fruit borne on racemes, panicles, or corymbs.   (10)
       
10 (9) Cauline leaves sessile, auriculate, sagittate, or amplexicaul; roots not fleshy, cylindric; fruit often winged apically   18 Thlaspi
+ Cauline leaves petiolate or sessile but not auriculate, sagittate, or amplexicaul; roots fleshy, fusiform or conical; fruit wingless or cristate apically.   (11)
       
11 (10) Plants 0.5-2 m tall; infructescence a large panicle; replum slender, not flattened; racemes ebracteate   42 Armoracia
+ Plants rarely to 0.3 m tall; infructescence a small raceme or corymb; replum broad, flattened at least basally; racemes often bracteate throughout.   (12)
       
12 (11) Cauline leaves sessile; fruit often cristate apically; fruit valves obscurely veined; seeds minutely reticulate   24 Dilophia
+ Cauline leaves petiolate; fruit not cristate apically; fruit valves with a prominent midvein; seeds papillate or foveolate   88 Taphrospermum
       
13 (8) Fruit didymous, breaking into 2, closed, 1-seeded halves.   (14)
+ Fruit not didymous, dehiscent and readily releasing seeds, or indehiscent.   (16)
       
14 (13) Fruit halves broadly winged, 8-40 mm; root fleshy, 1-15 cm in diameter   16 Megacarpaea
+ Fruit halves wingless, 1-2 mm; roots not fleshy, slender, much narrower.   (15)
       
15 (14) Dwarf plants rarely more than 3 cm tall; all or most fruit on solitary pedicels originating from basal rosette; leaves palmately veined; fruit halves smooth; stamens 6   17 Megadenia
+ Plants often more than 10 cm tall; fruit in racemes; leaves pinnately veined; fruit halves prominently reticulate to rugose, very rarely smooth; stamens 2 or 4   11 Coronopus
       
16 (13) Fruit 1-seeded, obscurely differentiated into halves, without a septum, indehiscent, with thickened, central, seed-bearing part; fruiting pedicels reflexed; petals yellow.   (17)
+ Fruit 2-seeded, well differentiated into halves, with a well-developed septum, dehiscent or rarely indehiscent, not thickened at center; fruiting pedicels not reflexed; petals white or pink (yellow in Lepidium perfoliatum).   (19)
       
17 (16) Fruit navicular, semiovoid to semiglobose, with strongly involute wing margins bearing 1 or 2 rows of tubercles; fruiting pedicels articulate at base   31 Tauscheria
+ Fruit oblong, elliptic, ovate, or pyriform, flat, wingless or wing margins neither involute nor tuberculate; fruiting pedicels not articulate.   (18)
       
18 (17) Fruit samaroid, more than 6 mm, wing margin not thickened   13 Isatis
+ Fruit achenelike, rarely to 5 mm, wing margin strongly thickened   14 Pachypterygium
       
19 (16) Fruit indehiscent; nectar glands confluent; style 0.7-2 mm   12 Cardaria
+ Fruit dehiscent; nectar glands separate teeth; style (ours) absent or to 0.8(-1) mm.   (20)
       
20 (19) Fruit somewhat 4-angled; leaves leathery; caudex thick, with petiolar remains of previous years; stamens 6   27 Stroganowia
+ Fruit flat, not angled; leaves herbaceous rarely leathery; caudex absent or slender, rarely thick and with petiolar remains of previous years; stamens 2, 4, or 6   10 Lepidium
       
21 (3) Fruit on solitary pedicels originating from basal rosette.   (22)
+ Fruit in racemes.   (25)
       
22 (21) Plants glabrous or with simple trichomes only; leaves entire, rarely toothed or pinnately lobed.   (23)
+ Plants with at least some branched trichomes; leaves subapically toothed.   (24)
       
23 (22) Fruit readily detached from pedicel; valve apex persistently united with replum; replum not flattened; valves marginal veins prominent; petals purple, pink, or blue   64 Solms-laubachia
+ Fruit not detached from pedicel; valve apex readily separated from replum at dehiscence; replum strongly flattened; valves marginal veins obscure or absent; petals white   44 Pegaeophyton
       
24 (22) Fruit ovate or lanceolate, not geocarpic; valves prominently veined; stigma 2-lobed   57 Desideria
+ Fruit oblong, geocarpic; valves obscurely veined; stigma entire   84 Pycnoplinthopsis
       
25 (21) Plants glabrous or with simple trichomes only.   (26)
+ Plants with at least some trichomes branched.   (36)
       
26 (25) Fruit indehiscent, with a woody or corky wall, sometimes breaking transversely into segments.   (27)
+ Fruit dehiscent, with a papery or membranous wall.   (30)
       
27 (26) Style slender, 1.5-3.5 mm, readily caducous from a basal articulation; replum strongly expanded laterally   33 Litwinowia
+ Style obsolete, or fruit attenuate into a conical, persistent, stylelike beak without a basal articulation; replum not expanded laterally.   (28)
       
28 (27) Cauline leaves petiolate, not auriculate; plants 0.5-2.5 m tall; fruit rounded at apex, strongly differentiated into a seedless basal segment and a larger, 1-seeded terminal segment   7 Crambe
+ Cauline leaves sessile, auriculate to amplexicaul; plants rarely to 0.5 m tall; fruit tapered to apex, unsegmented.   (29)
       
29 (28) Fruiting pedicels divaricate, not articulate at base; fruit 2-seeded, not constricted at middle; cotyledons spirolobal; petals white   35 Bunias
+ Fruiting pedicels reflexed, articulate at base; fruit 1-3-seeded, constricted between seeds; cotyledons incumbent; petals purplish or pink   82 Goldbachia
       
30 (26) At least some leaves dentate, lobed, or compound.   (31)
+ All leaves entire.   (33)
       
31 (30) Some or all cauline leaves auriculate or sagittate at base; seeds more than 50 per fruit; petals yellow   60 Rorippa
+ Cauline leaves not auriculate or sagittate at base; seeds less than 25 per fruit; petals white, pink, blue, or purple.   (32)
       
32 (31) Leaves trifoliolate, pinnately compound, pinnatisect, or rarely simple; racemes ebracteate, if bracteate, then basal bracts trifoliolate; fruiting pedicels glabrous or pubescent all around; petals white   26 Yinshania
+ Leaves palmately 3-5-lobed; racemes bracteate throughout; fruiting pedicels puberulent adaxially; petals purple, blue, or rarely pink   28 Lignariella
       
33 (30) Raceme bracteate throughout; fruit 1- or 2-seeded; valves with 3 rows of crests; filaments of median stamens appendaged   22 Hemilophia
+ Racemes ebracteate; fruit more than 2-seeded; valves without rows of crests; filaments of median stamens unappendaged.   (34)
       
34 (33) Fruit (0.8-)1-1.7 cm wide; valves with prominent midvein and lateral and marginal veins, apex persistently united with replum   64 Solms-laubachia
+ Fruit rarely to 0.3 cm wide; valves obscurely veined or only with a prominent midvein, apex readily separated from replum at dehiscence.   (35)
       
35 (34) Fruit slightly 4-angled, on a short gynophore; stems leafy; lowermost leaves cordate to ovate, often palmately veined   89 Eutrema
+ Fruit terete, sessile; stems leafless; lowermost leaves linear or oblong, pinnately veined   96 Braya
       
36 (25) Fruit indehiscent, often nutletlike, with woody or corky wall.   (37)
+ Fruit dehiscent, with papery wall.   (41)
       
37 (36) Plants with glandular trichomes, if eglandular then fruit strongly reflexed.   (38)
+ Plants eglandular, fruit erect to divaricate.   (39)
       
38 (37) Fruiting pedicels divaricate; trichomes simple and forked; seeds wingless, 1 or 2; cotyledons spirolobal; petals bright yellow   35 Bunias
+ Fruiting pedicels strongly reflexed; trichomes dendritic; seeds winged, often more than 2; cotyledons accumbent; petals yellowish white   67 Microstigma
       
39 (37) Cauline leaves sessile, auriculate or sagittate; style less than 1 mm; stigma entire; fruit readily detached from divaricate pedicels; petals yellow   34 Neslia
+ Cauline leaves petiolate, not auriculate or sagittate; style 1-7 mm; stigma deeply 2-lobed; fruit not readily detached from erect-ascending pedicels; petals white or pink.   (40)
       
40 (39) Fruit densely covered with silky trichomes 5-8 mm; style (2.5-)4-7 mm; petals pink   29 Lachnoloma
+ Fruit scabrous with minute trichomes less than 1 mm; style less than 2 mm; petals white   32 Euclidium
       
41 (36) Raceme bracteate throughout or at least along proximal half.   (42)
+ Raceme ebracteate.   (45)
       
42 (41) Fruit 1- or 2-seeded.   (43)
+ Fruit more than 4-seeded.   (44)
       
43 (42) Fruit valves with 3 rows of crests; fruiting pedicels straight or slightly curved, not forming a loop; leaves entire; filaments of median stamens appendaged   22 Hemilophia
+ Fruit valves without crests; fruiting pedicels often forming a loop; at least some leaves dentate or lobed near apex; filaments of median stamens unappendaged   23 Dipoma
       
44 (42) Fruit latiseptate; trichomes forked or stellate; cotyledons accumbent   41 Draba
+ Fruit terete; trichomes dendritic; cotyledons incumbent   100 Sophiopsis
       
45 (41) Leaves 1- or 2-pinnatisect or 3-5-foliolate.   (46)
+ Leaves entire or rarely dentate.   (48)
       
46 (45) Branched trichomes forked; inflorescence paniculate, ultimate branches flexuous   26 Yinshania
+ Branched trichomes dendritic; inflorescence non-flexuous racemes, if branched then fruit more than 10-seeded.   (47)
       
47 (46) Annuals or biennials; seeds mucilaginous when wetted; petals yellow   100 Sophiopsis
+ Pulvinate perennials; seeds not mucilaginous when wetted; petals white   101 Smelowskia
       
48 (45) Cauline leaves auriculate or sagittate at base; fruit obpyriform; valves connate at margin and hiding replum, beaklike apex caudate, extending into style   98 Camelina
+ Cauline leaves often neither auriculate nor sagittate at base; fruit variously shaped but not obpyriform; valve margins not connate, apex not extending into a caudate beak.   (49)
       
49 (48) Branched trichomes exclusively malpighiaceous.   (50)
+ Branched trichomes stellate or forked.   (51)
       
50 (49) Fruit 1- or 2- seeded; seeds orbicular; simple trichomes absent   38 Lobularia
+ Fruit more than 4-seeded; seeds oblong or ovate; simple trichomes present   41 Draba
       
51 (49) Branched trichomes exclusively stellate, sessile, appressed.   (52)
+ At least some of the branched trichomes simple or forked, stalked, not appressed.   (53)
       
52 (51) Seeds broadly winged (wing to 1 mm wide); petals deeply bifid; filaments not toothed or winged   37 Galitzkya
+ Seeds wingless or narrowly margined (margin to 0.1 mm wide); petals obtuse or emarginate; filaments often toothed or winged   36 Alyssum
       
53 (51) Cauline leaves absent.   (54)
+ Cauline leaves present.   (55)
       
54 (53) Style conical or subconical; fruit 5-10 mm wide; cotyledons incumbent, if accumbent then septum reduced to a rim   58 Eurycarpus
+ Style obsolete or cylindric; fruit rarely to 4 mm wide; cotyledons accumbent   41 Draba
       
55 (53) Fruit 1- or 2(-4)-seeded; stamens often toothed or winged   36 Alyssum
+ Fruit more than 4-seeded; stamens neither toothed nor winged.   (56)
       
56 (55) Fruit torulose; cotyledons incumbent   96 Braya
+ Fruit not torulose, smooth; cotyledons accumbent.   (57)
       
57 (56) Annuals or biennials; petals deeply bifid; seed narrowly margined   39 Berteroa
+ Perennials, or if annuals then styles obsolete; petals obtuse or rarely emarginate, sometimes absent; seeds not margined.   (58)
       
58 (57) Plants stoloniferous; filament bases broadly dilated   40 Coelonema
+ Plants cespitose, perennials or rarely annuals; filament bases not dilated   41 Draba
       
59 (1) Fruit borne on solitary pedicels originating from basal rosette.   (60)
+ Fruit borne on racemes, corymbs, or panicles.   (67)
       
60 (59) Plants with glandular trichomes; fruit lomentaceous, breaking into 1-seeded corky units   68 Chorispora
+ Plants without glandular trichomes; fruit dehiscent, not breaking into units.   (61)
       
61 (60) Stigma conical, prominently 2-lobed, lobes decurrent; seeds often broadly winged   72 Leiospora
+ Stigma capitate, entire or slightly 2-lobed and lobes not decurrent; seeds wingless.   (62)
       
62 (61) Plants with branched trichomes.   (63)
+ Plants glabrous or with simple trichomes only.   (65)
       
63 (62) Leaves entire; plants ca. 2 cm tall   86 Baimashania
+ At least some leaves coarsely dentate; plants much coarser.   (64)
       
64 (63) Fruit strongly flattened; valves prominently veined, apex persistently united with replum; style obsolete; calyx deciduous before fruit maturity; cotyledons accumbent   57 Desideria
+ Fruit terete; valves obscurely veined, readily separated from replum at dehiscence; style distinct; calyx persistent at fruit base; cotyledons incumbent   84 Pycnoplinthopsis
       
65 (62) Fruit valves obscurely veined; replum flattened; fruiting pedicels persistent next season, puberulent along 1 line   44 Pegaeophyton
+ Fruit valves prominently veined; replum terete; fruiting pedicels neither persistent nor puberulent along 1 line.   (66)
       
66 (65) Style obsolete; valve apex persistently united with replum; cotyledons accumbent   64 Solms-laubachia
+ Style distinct, to 1 mm; valve apex readily separated from replum at dehiscence; cotyledons incumbent   85 Pycnoplinthus
       
67 (59) Glandular trichomes present.   (68)
+ Glandular trichomes absent.   (77)
       
68 (67) Branched trichomes present.   (69)
+ Branched trichomes absent.   (71)
       
69 (68) Fruit and fruiting pedicels strongly reflexed; cotyledons accumbent   67 Microstigma
+ Fruit and fruiting pedicels divaricate to ascending; cotyledons incumbent.   (70)
       
70 (69) Fruit valves with a prominent midvein; branched trichomes forked; stigma lobes decurrent; glands on uniseriate, few-celled stalks; median filaments not united   75 Hesperis
+ Fruit valves not or obscurely veined; branched trichomes dendritic; stigma lobes not decurrent; glands on multiseriate, multicellular stalks; median filaments united in pairs   81 Oreoloma
       
71 (68) Stigma slightly lobed, lobes neither decurrent nor connivent; median filaments united in pairs or strongly flattened; lateral sepals not saccate   63 Dontostemon
+ Stigma prominently lobed; lobes decurrent, connivent; median filaments neither united nor strongly flattened; lateral sepals saccate.   (72)
       
72 (71) Fruit terete; seeds wingless.   (73)
+ Fruit (at least distal ones) flattened; seeds winged.   (74)
       
73 (72) Fruit breaking transversely into 1-seeded corky units; style 1-9 mm; cotyledons accumbent   68 Chorispora
+ Fruit dehiscent, not breaking into corky units; style obsolete; cotyledons incumbent   75 Hesperis
       
74 (72) Fruit dimorphic, lowermost indehiscent, corky, and terete, uppermost dehiscent and flattened; replum rounded   69 Diptychocarpus
+ Fruit of 1 kind, dehiscent, flattened, terete; replum flattened.   (75)
       
75 (74) Cauline leaves absent, rarely few; fruit (2-)2.5-7 mm wide; seeds broadly winged, wing 0.5-3 mm wide   70 Parrya
+ Cauline leaves several; fruit 1.5-2 mm wide; seeds narrowly winged, wing 0.1-0.4 mm wide.   (76)
       
76 (75) Basal leaves not rosulate; stigma capitate, lobes slightly decurrent   76 Clausia
+ Basal leaves rosulate; stigma linear, lobes strongly decurrent   77 Pseudoclausia
       
77 (67) Plants glabrous or with simple trichomes only.   (78)
+ Plants with branched trichomes.   (121)
       
78 (77) Fruit indehiscent, sometimes breaking transversely into corky, 1-seeded units.   (79)
+ Fruit dehiscent.   (83)
       
79 (78) Cauline leaves auriculate or amplexicaul; style obsolete or rarely to 2 mm; lateral sepals not saccate.   (80)
+ Cauline leaves not auriculate or amplexicaul; style distinct, 5-50 mm; lateral sepals often strongly saccate.   (81)
       
80 (79) Fruit a samara, strongly angustiseptate, 1-seeded; petals yellow   13 Isatis
+ Fruit nutletlike, terete or angled, 1-3 seeded, if 2- or 3- seeded then breaking transversely into 1-seeded units; petals purple or pink   82 Goldbachia
       
81 (79) Fruit 1-seeded, S-shaped, with a slender, stipelike base; fruiting pedicels articulate at base   30 Spirorhynchus
+ Fruit more than 2-seeded, linear, lanceolate, or ovoid, sessile or nearly so; fruiting pedicels not articulate at base.   (82)
       
82 (81) Stigma capitate, entire or slightly lobed; cotyledons conduplicate; plants annual or biennial; petal veins darker in color than rest of petal   6 Raphanus
+ Stigma conical, with prominently decurrent and connivent lobes; cotyledons accumbent; plants perennial; petals uniform in color   68 Chorispora
       
83 (78) Raceme bracteate throughout or rarely only along proximal half.   (84)
+ Raceme ebracteate.   (92)
       
84 (83) Cauline leaves palmately veined.   (85)
+ Cauline leaves pinnately veined.   (88)
       
85 (84) Fruit flattened, valves coiled during dehiscence   43 Cardamine
+ Fruit terete or 4-angled, valves not coiled during dehiscence.   (86)
       
86 (85) Leaves with distinct apiculate callosities terminating ultimate veins   89 Eutrema
+ Leaves without apiculate callosities terminating ultimate veins.   (87)
       
87 (86) Taproot fleshy, fusiform; stigma 2-lobed; fruiting pedicels glabrous; cotyledons accumbent; median staminal filaments 1-toothed; petals white   25 Platycraspedum
+ Taproot not fleshy, slender; stigma entire; fruiting pedicels papillate adaxially; cotyledons incumbent; petals purple   28 Lignariella
       
88 (84) Fruit 90-200-seeded; plants annual   60 Rorippa
+ Fruit 4-30-seeded; plants biennial or perennial with caudices, rhizomes, or fleshy taproots.   (89)
       
89 (88) Fruit valves with prominent midvein and marginal veins, apex persistently united with replum; at least some leaves dentate; fruit 14-26-seeded; style obsolete   57 Desideria
+ Fruit valves usually obscurely veined, readily separated from replum at dehiscence; leaves entire; fruit 4-12-seeded; style more than 0.5 mm.   (90)
       
90 (89) Replum flattened; roots fleshy; plants without rhizomes or caudices; basal leaves not persistent   88 Taphrospermum
+ Replum terete; roots not fleshy; plants with rhizomes or caudices; basal leaves or petioles often persistent.   (91)
       
91 (90) Fruit not torulose; cotyledons incumbent   45 Phaeonychium
+ Fruit torulose; cotyledons accumbent   62 Lepidostemon
       
92 (83) Fruit angustiseptate   18 Thlaspi
+ Fruit latiseptate, terete, or angled.   (93)
       
93 (92) Stigma conical or cylindric, with prominent, decurrent lobes.   (94)
+ Stigma capitate, entire, or if lobed then lobes not decurrent.   (98)
       
94 (93) Fruit latiseptate; seeds broadly winged; cauline leaves absent or rarely few.   (95)
+ Fruit terete or 4-angled; seeds wingless; cauline leaves several.   (96)
       
95 (94) Style distinct, 0.5-6 mm; valve apex readily separated at dehiscence from replum   70 Parrya
+ Style absent; valve apex persistently united with replum   72 Leiospora
       
96 (94) Fruit not torulose, segmented, with stylelike, ensiform, 5-veined, terminal segment   5 Eruca
+ Fruit torulose, unsegmented, with a distinct style 1-3 mm.   (97)
       
97 (96) Leaves pectinate-pinnatifid, terminal lobe about as large as lateral ones; cotyledons accumbent   63 Dontostemon
+ Leaves simple or pinnatisect, terminal lobe considerably larger than lateral ones; cotyledons conduplicate   8 Orychophragmus
       
98 (93) Cotyledons conduplicate; fruit often segmented, usually with a beaklike terminal segment; seeds globose, rarely ovoid, ellipsoid, or oblong.   (99)
+ Cotyledons accumbent or incumbent; fruit unsegmented, not beaked at apex; seeds variously shaped but never globose.   (102)
       
99 (98) Fruit flattened; seeds biseriate   4 Diplotaxis
+ Fruit terete or 4-angled; seeds uniseriate.   (100)
       
100 (99) Fruit valves prominently 3-7-veined   3 Sinapis
+ Fruit valves obscurely veined or only midvein prominent.   (101)
       
101 (100) Stigma entire; seeds globose; fruit often segmented, without style or with stylelike beak; petals yellow   2 Brassica
+ Stigma distinctly 2-lobed, lobes subdecurrent; seeds oblong; fruit not segmented, with a distinct style; petals purple, lavender, or white   8 Orychophragmus
       
102 (98) Fruit valves without a midvein, always coiled during dehiscence; replum strongly flattened or winged   43 Cardamine
+ Fruit valves with a prominent midvein or sometimes obscurely veined, never coiled during or after dehiscence; replum terete or rarely slightly flattened (a few Dontostemon).   (103)
       
103 (102) All leaves palmately veined, undivided and reniform, cordate, deltoid, or broadly ovate.   (104)
+ Leaves pinnately veined, pinnately divided, if undivided then shapes different from above.   (108)
       
104 (103) Leaf margin with distinct, mucronate callosities terminating ultimate veins; fruit valves obscurely veined.   (105)
+ Leaf margin without mucronate callosities; fruit valves prominently veined.   (106)
       
105 (104) Annuals; fruit more than 10-seeded; cotyledons accumbent   47 Neomartinella
+ Rhizomatous perennials; fruit less than 10 seeded; cotyledons incumbent   89 Eutrema
       
106 (104) Leaves flabellate, obovate, or spatulate; fruit latiseptate, valve apex persistently united with replum; cotyledons accumbent   57 Desideria
+ Leaves reniform, ovate, or deltoid; fruit terete or 4-angled, valve apex readily separated from replum at dehiscence; cotyledons incumbent.   (107)
       
107 (106) Biennials; fruit torulose, valves 3-veined; seeds longitudinally striate   87 Alliaria
+ Perennials; fruit not torulose, valves 1-veined; seeds reticulate   89 Eutrema
       
108 (103) At least lowermost cauline leaves pinnatifid, pinnatisect, or pinnately compound.   (109)
+ Leaves entire or dentate.   (113)
       
109 (108) Fruit valves not veined or obscurely veined, if with a distinct midvein then seeds biseriate and colliculate.   (110)
+ Fruit valves with prominent midvein and marginal veins.   (111)
       
110 (109) Leaves pinnatisect or lyrate; stems solid, not rooting from lowermost nodes; seeds foveolate or colliculate; petals yellow   60 Rorippa
+ Leaves pinnately compound; stems hollow, rooting from lowermost nodes; seeds coarsely reticulate; petals white   61 Nasturtium
       
111 (109) Upper cauline leaves auriculate; stems distinctly angular   46 Barbarea
+ Upper cauline leaves not auriculate; stems usually terete.   (112)
       
112 (111) Fruit flattened; stigma entire; cotyledons accumbent; petals white or pink   49 Sisymbriopsis
+ Fruit terete; stigma 2-lobed; cotyledons incumbent; petals yellow   90 Sisymbrium
       
113 (108) Cauline leaves absent.   (114)
+ Cauline leaves present.   (115)
       
114 (113) Fruit lanceolate, latiseptate; basal leaves setose-ciliate   45 Phaeonychium
+ Fruit ovoid, terete; basal leaves not setose-ciliate   96 Braya
       
115 (113) Seeds foveolate or colliculate, 60-110 per fruit; petals yellow; cotyledons accumbent   60 Rorippa
+ Seeds reticulate, fewer per fruit, (to 96 in Thellungiella salsuginea); petals white, lavender, or purple (yellow in Christolea); cotyledons incumbent (accumbent in some Dontostemon).   (116)
       
116 (115) Annuals.   (117)
+ Perennials.   (119)
       
117 (116) Raceme rachis with minute, twisted trichomes; plants not glaucous; cauline leaves not auriculate or amplexicaul; filaments of median stamens united in pairs   63 Dontostemon
+ Raceme rachis glabrous; plants glaucous; cauline leaves auriculate or amplexicaul (except for Thellungiella parvula); filaments of median stamens free.   (118)
       
118 (117) Fruit 3-10 cm, valves with an obscure midvein; seeds 1.5-2 mm   9 Conringia
+ Fruit 0.2-2 cm, valves with a prominent midvein; seeds 0.4-0.8 mm   97 Thellungiella
       
119 (116) At least middle and lower cauline leaves petiolate, dentate; seeds transversely oriented in fruit; petals yellow   56 Christolea
+ Leaves sessile, entire; seeds longitudinally oriented in fruit; petals white or purple.   (120)
       
120 (119) Seeds 20-60 per fruit; fruit terete or latiseptate; median staminal filaments united in pairs   63 Dontostemon
+ Seeds 2-10 per fruit; fruit slightly 4-angled; median staminal filaments free   89 Eutrema
       
121 (77) Cauline leaves 1- or 2-pinnatisect or narrowly trifid.   (122)
+ Cauline leaves undivided, lyrate, or rarely pinnatifid.   (129)
       
122 (121) Fruit angustiseptate   20 Hedinia
+ Fruit terete or latiseptate.   (123)
       
123 (122) Fruit latiseptate; filaments of median stamens united in pairs.   (124)
+ Fruit terete or slightly 4-angled; filaments of median stamens free.   (125)
       
124 (123) Fruit tardily dehiscent, not torulose; leaf segments filiform; stigma conical; seeds wingless   73 Leptaleum
+ Fruit readily dehiscent, torulose; ultimate leaf segment oblong; stigma capitate; seeds narrowly winged distally   95 Synstemon
       
125 (123) Infructescence rachis flexuous; uppermost leaves narrowly trifid   91 Ianhedgea
+ Infructescence rachis straight; uppermost leaves pinnatisect, rarely pinnatifid.   (126)
       
126 (125) Dendritic trichomes absent; valves obscurely veined or not veined   102 Sinosophiopsis
+ Dendritic trichomes present; valves often prominently veined.   (127)
       
127 (126) Pulvinate perennials; petals white or rarely pale yellow; style 0.5-1.5 mm; seeds not mucilaginous when wetted   101 Smelowskia
+ Annuals or biennials; petals yellow; style obsolete or rarely to 0.5 mm; seeds mucilaginous when wetted.   (128)
       
128 (127) Fruit linear, 20-40-seeded, more than 1 cm, torulose   99 Descurainia
+ Fruit obovoid to ellipsoid, rarely ellipsoid-linear, 4-16-seeded, rarely to 1 cm, not torulose   100 Sophiopsis
       
129 (121) Fruit subapically 4-horned   65 Tetracme
+ Fruit not horned.   (130)
       
130 (129) Stigma conical, with strongly decurrent, connivent or united lobes.   (131)
+ Stigma capitate, entire or slightly lobed, rarely with divergent lobes.   (134)
       
131 (130) Fruit latiseptate; seeds broadly winged; trichomes stellate and/or dendritic; cotyledons accumbent.   (132)
+ Fruit terete or 4-angled (latiseptate in Malcolmia hispida and M. karelinii); seeds wingless; trichomes simple and forked; cotyledons incumbent.   (133)
       
132 (131) Perennials; fruit 6-11 cm × 1.5-2.7 mm; petals linear, circinately involute   66 Matthiola
+ Annuals; fruit 3-5 cm × 3-6 mm; petals oblong-obovate to narrowly oblanceolate, flat   71 Cithareloma
       
133 (131) Annuals; lateral sepals not saccate   74 Malcolmia
+ Perennials or biennials; lateral sepals saccate   75 Hesperis
       
134 (130) At least fruit, pedicels, and distal portions of stem exclusively or predominately with malpighiaceous trichomes and without simple trichomes ones.   (135)
+ Malpighiaceous trichomes absent from distal portions of plant.   (137)
       
135 (134) Fruit indehiscent, breaking transversely into 1-seeded segments; septum absent; lowermost parts of plant with stalked trichomes   79 Cryptospora
+ Fruit dehiscent; septum present; all parts of plant with sessile, appressed trichomes.   (136)
       
136 (135) Cotyledons accumbent; petals white becoming purple; anthers of median stamens 1-lobed   78 Atelanthera
+ Cotyledons incumbent; petals yellow or orange; anthers of median stamens 2-lobed   83 Erysimum
       
137 (134) Fruit (3-)3.5-7(-10) mm wide.   (138)
+ Fruit to 2(-2.5) mm wide.   (142)
       
138 (137) Fruit on a recurved gynophore 3-5 mm; stamens long exserted, 7-10 mm   1 Macropodium
+ Fruit sessile; stamens included or slightly exserted, less than 7 mm.   (139)
       
139 (138) Racemes bracteate throughout   57 Desideria
+ Racemes ebracteate.   (140)
       
140 (139) Fruit strongly reflexed, 4-angled, indehiscent; seeds winged   67 Microstigma
+ Fruit erect to divaricate, latiseptate, dehiscent; seeds wingless.   (141)
       
141 (140) Fruit margins strongly indurated, bases of valves and replum persistently united; petals white; filaments of median stamens winged   62 Lepidostemon
+ Fruit margins not indurated, bases of valves and replum readily separated at dehiscence; petals purplish; median stamens slender   58 Eurycarpus
       
142 (137) Fruit indehiscent, breaking transversely into 1-seeded segments; filaments of median stamens united to middle or nearly to apex   80 Sterigmostemum
+ Fruit dehiscent; filaments of median stamens free, rarely united only at base (Synstemon petrovii).   (143)
       
143 (142) Seeds winged at least distally.   (144)
+ Seeds wingless.   (146)
       
144 (143) Cotyledons incumbent; filaments of median stamens united at base; petal claws pilose   95 Synstemon
+ Cotyledons accumbent; filaments of median stamens free; petal claws glabrous.   (145)
       
145 (144) Cauline leaves auriculate at base, or absent; valves obscurely veined or only midvein distinct; fruit elliptic-linear in cross section   48 Arabis
+ Cauline leaves petiolate; valves with prominent midvein and 2 marginal veins and conspicuous lateral ones; fruit rectangular in cross section   49 Sisymbriopsis
       
146 (143) Fruit latiseptate.   (147)
+ Fruit terete or 4-angled.   (158)
       
147 (146) Fruiting stems leafless.   (148)
+ Fruiting stems leafy.   (152)
       
148 (147) Annuals or biennials.   (149)
+ Perennials with well-developed caudex.   (150)
       
149 (148) Fruit valve obscurely veined; seeds biseriate   41 Draba
+ Fruit valve with a prominent midvein; seeds uniseriate   55 Drabopsis
       
150 (148) Seeds biseriate; petals yellow   41 Draba
+ Seeds uniseriate; petals white or pink.   (151)
       
151 (150) Plants robust, not pulvinate, 8-30 cm tall; cotyledons incumbent; petals 6.5-10 mm   45 Phaeonychium
+ Plants slender, pulvinate, to 2 cm tall; cotyledons accumbent; petals 3-4 mm   86 Baimashania
       
152 (147) Racemes bracteate throughout or at least along proximal part.   (153)
+ Racemes ebracteate.   (155)
       
153 (152) Cotyledons accumbent; fruits often twisted; seeds biseriate   41 Draba
+ Cotyledons incumbent; fruits not twisted; seeds uniseriate.   (154)
       
154 (153) Fruit linear; seeds uniseriate   45 Phaeonychium
+ Fruit lanceolate or elliptic; seeds biseriate   92 Aphragmus
       
155 (152) Plants exclusively with appressed, stellate trichomes; lateral sepals subsaccate; replum constricted between seeds; seeds 6-12 per fruit   54 Stevenia
+ Plants with more than 1 kind of trichomes; lateral sepals often not saccate; replum not constricted between seeds; seeds (16-)20-80 per fruit.   (156)
       
156 (155) Seeds biseriate; fruit narrowly oblong, lanceolate, lanceolate-linear, or rarely linear, often slightly to strongly contorted   41 Draba
+ Seeds uniseriate; fruit narrowly linear, not contorted.   (157)
       
157 (156) Cauline leaves sessile, auriculate, sagittate, or amplexicaul at base; at least some trichomes stellate   48 Arabis
+ Cauline leaves petiolate, not auriculate at base; trichomes simple and forked   50 Arabidopsis
       
158 (146) Cauline leaves sessile, auriculate, sagittate, or amplexicaul at base.   (159)
+ Cauline leaves petiolate or narrowed to a petiolelike base, never auriculate, or fruiting stems leafless.   (162)
       
159 (158) Fruit nearly 4-angled, appressed to rachis; plants glaucous distally; cotyledons accumbent   59 Turritis
+ Fruit terete, variously oriented, if appressed to stem then racemes bracteate and/or fruit pubescent; plants not glaucous distally; cotyledons incumbent.   (160)
       
160 (159) Branched trichomes sessile, all stellate   53 Pseudoarabidopsis
+ Branched trichomes forked, submalpighiaceous, subdendritic, and stellate, often more than 1 type present.   (161)
       
161 (160) Fruit with submalpighiaceous trichomes; petals yellow or rarely creamy white; racemes ebracteate   52 Olimarabidopsis
+ Fruit glabrous or with stellate or forked trichomes; petals purple, pink, or white; racemes often bracteate at least basally   51 Crucihimalaya
       
162 (158) Fruit pubescent.   (163)
+ Fruit glabrous.   (165)
       
163 (162) Plants with exclusively stalked, stellate trichomes; filaments of median stamens strongly flattened or winged at base   93 Berteroella
+ Plants with a mixture of simple, forked, stellate, or submalpighiaceous trichomes; filaments of median stamens slender.   (164)
       
164 (163) Fruit linear, sometimes curved or twisted; stems often subdecumbent; plants annual, if perennial then racemes bracteate at least basally   94 Neotorularia
+ Fruit oblong, ovoid, or rarely linear; stems erect; plants perennial, with ebracteate racemes   96 Braya
       
165 (162) Fruiting stems leafy.   (166)
+ Fruiting stems leafless.   (167)
       
166 (165) Plants with exclusively simple and forked trichomes; upper cauline leaves entire; seeds 0.3-0.5 mm   50 Arabidopsis
+ Plants with some stellate and/or dendritic trichomes; at least some upper cauline leaves dentate, denticulate, or lobed; seeds larger   51 Crucihimalaya
       
167 (165) Annuals; fruit linear; fruiting pedicels nearly as wide as fruit   94 Neotorularia
+ Perennials; fruit ovoid, oblong, or rarely linear; fruiting pedicels narrower than fruit   96 Braya

List of Keys

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  • Comments

    Ihsan A. Al-Shehbaz
    12/22/2004
    Taxon name changes since publication:

    • Coronopus integrifolius: change to Lepidium englerianum (Muschler) Al-Shehbaz

    • Coronopus didymus: change to Lepidium didymum Linnaeus

    • Cardaria draba subsp. draba: change to Lepidium draba Linnaeus

    • Cardaria draba subsp. chalapensis: change to Lepidium chalepense Linnaeus

    • Cardaria pubescens: change to Lepidium appelianum Al-Shehbaz

    • Thlaspi perfoliatum: change to Microthlaspi perfoliatum (Linnaeus) F. K. Meyer

    • Thlaspi andersonii: change to Noccaea andersonii (J. D. Hooker & Thomson) Al-Shehbaz

    • Thlaspi flagelliferum: change to Noccaea flagillferum (O. E. Schulz) Al-Shehbaz

    • Thlaspi yunnanense: change to Noccaea yunnanense (Franchet) Al-Shehbaz

    • Hedinia tibetica: change to Smelowskia tibetica (Thomson) Lipsky

    • Stroganowia brachyota: change to Lepidium brachyotum (Karelin & Kirilov) Al-Shehbaz

    • Coelonema draboides: change to Draba draboides (Maximowicz) Al-Shehbaz

    • Draba serpens: change to Hemilophia serpens (O. E. Schulz) Al-Shehbaz

    • Solms-laubachia minor: change to Solms-laubachia zhongdianensis J. P. Yue, Al-Shehbaz & H. Sun

    • Neotorularia humilis: change to Braya humilis (C. A. Meyer) B. L. Robinson

    • Neotorularia brachycarpa: change to Braya parvia (Z. X. An) Al-Shehbaz

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