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A synopsis of the South American Lepidium (Brassicaceae)

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A synopsis of the South American Lepidium (Brassicaceae)

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Fifty native and 12 naturalized species of Lepidium grow in South America, and a key to all 62 species is presented. Three new species from Argentina (L. hickenii, L. pedersenii, and L. santacruzensis) and two from Peru (L. cuzcoensis and L. werffii) are described, illustrated, and their relationships to nearest relatives are discussed. The new combination Lepidium crassius is proposed. Furthermore, 19 taxa (Coronopus didymus var. macrocarpus, C. didymus var. procumbens, L. abrotanifolium var. stein-mannii, L. affine, L. argentinum, L. auriculatum, L. bonariense var. pseudovirginicum, L. calycinum var. integrifolium, L. costaricense, L. cumingianum var. orbiculatum, L. cumingianum subsp. bertero-anum, L. depressum, L. myrianthum, L. neglectum, L. parodii, L. pubescens var. fallax, L. subvagina-tum, L. virginicum subsp. centrali-americanum, Thlaspi campestre) are lectotypified and 17 others (Coronopus leptocarpus, C. leptocarpus var. microcarpus, L. boelckii, L. bonariense var. gayi, L. bonariense var. pseudovirginicum, L. brevicaule, L. calycinum, L. danielsii, L. demissum, L. kalenbor-nii, L. morrisonii, L. peruvianum, L. philippianum var. boliviense, L. raimondii, L. scabrifructum, L. spicatum var. caylx-persistente, L. subvaginatum) are reduced to synonymy. L. depressum and L. rah-meri are new records for Argentina.
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Darwiniana
ISSN: 0011-6793
sdenham@darwin.edu.ar
Instituto de Botánica Darwinion
Argentina
Al-Shehbaz, Ihsan A.
A synopsis of the South American Lepidium (Brassicaceae)
Darwiniana, vol. 48, núm. 2, agosto-diciembre, 2010, pp. 141-167
Instituto de Botánica Darwinion
Buenos Aires, Argentina
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INTRODUCTION
The limits of Lepidium (Brassicaceae) were
initially expanded by Al-Shehbaz et al. (2002)
to include Cardaria Desv., Coronopus Zinn,
and Stroganowia Kar. & Kir., and more recently
by Al-Shehbaz and Mummenhoff (2010) to
include Stubendorffia Schrenk ex Fisch. and
141
DARWINIANA 48(2): 141-167. 2010 ISSN 0011-6793
A SYNOPSIS OF THE SOUTH AMERICAN LEPIDIUM (BRASSICACEAE)
Ihsan A. Al-Shehbaz
Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A.; ihsan.al-shehbaz@mobot.org
Original recibido el 2 de septiembre de 2010, aceptado el 30 de noviembre de 2010.
Abstract. Al-Shehbaz, I. A. 2010. A synopsis of the South American Lepidium (Brassicaceae). Darwiniana 48(2):
141-167.
Fifty native and 12 naturalized species of Lepidium grow in South America, and a key to all 62 spe-
cies is presented. Three new species from Argentina (L. hickenii, L. pedersenii, and L. santacruzensis)
and two from Peru (L. cuzcoensis and L. werffii) are described, illustrated, and their relationships to
nearest relatives are discussed. The new combination Lepidium crassius is proposed. Furthermore, 19
taxa (Coronopus didymus var. macrocarpus, C. didymus var. procumbens, L. abrotanifolium var. stein-
mannii, L. affine, L. argentinum, L. auriculatum, L. bonariense var. pseudovirginicum, L. calycinum
var. integrifolium, L. costaricense, L. cumingianum var. orbiculatum, L. cumingianum subsp. bertero-
anum, L. depressum, L. myrianthum, L. neglectum, L. parodii, L. pubescens var. fallax, L. subvagina-
tum, L. virginicum subsp. centrali-americanum, Thlaspi campestre) are lectotypified and 17 others
(Coronopus leptocarpus, C. leptocarpus var. microcarpus, L. boelckii, L. bonariense var. gayi, L.
bonariense var. pseudovirginicum, L. brevicaule, L. calycinum, L. danielsii, L. demissum, L. kalenbor-
nii, L. morrisonii, L. peruvianum, L. philippianum var. boliviense, L. raimondii, L. scabrifructum, L.
spicatum var. caylx-persistente, L. subvaginatum) are reduced to synonymy. L. depressum and L. rah-
meri are new records for Argentina.
Keywords. Brassicaceae, Lepidium, South America, taxonomy.
Cincuenta especies nativas y 12 naturalizadas de Lepidium crecen en América del Sur, se presenta
una clave para las 62 especies. Se describen e ilustran tres nuevas especies de Argentina (L. hickenii,
L. pedersenii y L. santacruzensis) y dos para Perú (L. cuzcoensis y L. werffii), y se discuten sus rela-
ciones con sus especies más afines. Se propone Lepidium crassius comb. nov., se designa lectotipo
para 19 binomios (Coronopus didymus var. macrocarpus, C. didymus var. procumbens, L. abrotanifo-
lium var. steinmannii, L. affine, L. argentinum, L. auriculatum, L. bonariense var. pseudovirginicum,
L. calycinum var. integrifolium, L. costaricense, L. cumingianum var. orbiculatum, L. cumingianum
subsp. berteroanum, L. depressum, L. myrianthum,L. neglectum, L. parodii, L. pubescens var. fallax,
L. subvaginatum, L. virginicum subsp. centrali-americanum, Thlaspi campestre), y se reducen a sinó-
nimos otros 17 nombres (Coronopus leptocarpus, C. leptocarpus var. microcarpus, L. boelckii, L.
bonariense var. gayi, L. bonariense var. pseudovirginicum, L. brevicaule, L. calycinum, L. danielsii, L.
demissum, L. kalenbornii, L. morrisonii, L. peruvianum, L. philippianum var. boliviense, L. raimondii,
L. scabrifructum,L. spicatum var. caylx-persistente, L. subvaginatum). L. depressum y L. rahmeri son
nuevos registros para la Argentina.
Palabras clave. Brassicaceae, Lepidium, Sudamérica, taxonomía.
Resumen. Al-Shehbaz, I. A. 2010. Sinopsis de las especies sudamericanas de Lepidium (Brassicaceae). Darwiniana
48(2): 141-167.
142
DARWINIANA 48(2) 141-167. 2010
Winklera Regel, based on extensive molecular
studies by Brüggemann (2000) and Mummen-
hoff et al. (2001, 2009, and references therein).
Lepidium is now represented by native species
on all continents except Antarctica, and as
currently recognized, the genus has 230 spe-
cies.
During work on the accounts of Brassicace-
ae (Cruciferae) for Catálogo de las Plantas
Vasculares del Cono Sur (Al-Shehbaz, 2008)
and for Flora de la República Argentina, seve-
ral new species of Lepidium L. were discove-
red. Furthermore, an examination of the types
of nearly all South American taxa of the genus
necessitated several nomenclatural adjustments
and lectotypifications. As a result, it became
clear that a synopsis of the entire genus for the
continent is needed, and it is provided herein
along with a key to all native and naturalized
species.
MATERIALS AND METHODS
The present study is based on loans of over
2,500 specimens (including types) of South
American Lepidium from most of the major her-
baria. In addition, databases and digital images
on-line were consulted, as well as additional
images were directly requested (see acknowled-
gements).
RESULTS AND TAXONOMIC TREATMENT
Fifty native and 12 naturalized species grow in
South America. The majority of native species
grow in Argentina (24 spp., 10 endemic), followed
by Chile (18 spp., seven endemic), Bolivia (12
spp., four endemic), and Peru (11 spp., four ende-
mic). The following key deals with all 62 South
American species of Lepidium.
1. At least some cauline leaves auriculate, sagittate, or amplexicau . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1. Cauline leaves not auriculate, petiolate or sessile, sometimes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22
2(1). Fruits indehiscent; plants perennial, rhizomatous; fruiting racemes hardly elongated . . . . . . . . . . . . . . . . . . . . .3
2. Fruits dehiscent; plants annual or biennial, rarely perennial with caudex; fruiting racemes strongly elongated . . .5
3(2). Fruits cordate to reniform, flattened; valves reticulately veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. draba
3. Fruits globose, subglobose, or ovoid, inflated; valves not reticulately veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
4(3). Fruits pubescent, (2-)3-4(-5) mm in diam.; style 0.7-1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. appelianum
4. Fruits glabrous, (3.5-)4-6.2(-7) mm in diam.; style (0.8-)1.2-2(-2.3) mm . . . . . . . . . . . . . . . . . . . . . .L. chalepense
5(2). Petals yellow; uppermost leaves deeply cordate-amplexicaul at base, entire . . . . . . . . . . . . . . . . .L. perfoliatum
5. Petals white, rudimentary, or absent; uppermost leaves auriculate or sagittate, entire, dentate, pinnatifid,
or pinnatisect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6(5). Stamens 6; fruit wings united with basal part of style; naturalized weeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
6. Stamens 2, rarely 4; fruit wings not united with basal part of style; native species . . . . . . . . . . . . . . . . . . . . . . . . .8
7(6). Annuals or biennials; fruit valves papillate; style 0.2-0.5(-0.7) mm, included in apical notch . . . . .L. campestre
7. Perennials with caudex; fruit valves often not papillate; style (0.6-)1-1.5 mm, well exserted
from apical notch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. heterophyllum
8(6). Annuals or rarely biennial without caudex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
8. Perennials with slender to well-developed caudex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
9(8). Stems prostrate, several from base, branched throughout; style as long as apical notch of fruit;
seeds wingless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. santacruzensis
9. Stems erect, often simple at base, branched mainly above; style shorter that apical notch of fruit; seeds winged or
margined at least apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
10(9). All cauline leaves 1-3-pinnatisect or -pinnatifid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. auriculatum
10. At least some cauline leaves entire, dentate, serrate, or incised . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11
11(10). Fruits wider than long; petals absent; North Chile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. johnstonii
11. Fruits longer than broad, rarely as long as wide; petals present; Argentina, Patagonian Chile, Paraguay . . . . . .12
12(11). Uppermost leaves linear to linear-lanceolate; basal leaves pinnatisect; fruit 2-2.6 mm; sepals caducous or
rarely persistent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. spicatum
12. Uppermost leaves oblong, ovate, or lanceolate; basal leaves dentate, serrate, or serrulate; fruit 2.5-3.5 mm; sepals
persistent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
Key to Lepidium species
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
143
13(12). Stems 1.5-4.5 dm; middle and upper leaves subentire or serrulate; fruiting pedicels strigillose
adaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. pedersenii
13. Stems 0.5-1.2(-1.5) dm; middle and upper cauline leaves incised; fruits 2.5-3.2 mm; fruiting pedicels puberulent
all around . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. tandilense
14(8). Stamens 4; petals 2-3 mm; apical notch of fruit absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. crassius
14. Stamens 2; petals 0.3-1.5(-2) mm; apical notch of fruit 0.1-1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15
15(14). Fruits wingless, wider than long; style 0.35-0.5 mm; petals 1.5-2 mm . . . . . . . . . . . . . . . . . . .L. spathulatum
15. Fruits winged, longer than wide, rarely length and width subequal; style obsolete or to 0.2 mm; petals 0.3-1.5 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16
16(15). Middle and upper cauline leaves pinnatifid or pinnatisect, rarely laciniate, with 3-6(-8) lobes
on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. bipinnatifidum
16. Middle and upper cauline leaves entire or toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17
17(16). Fruits 2.2-3 × 2-2.5 mm; fruiting pedicels 1.5-2(-3) mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18
17. Fruits 3-5.3 × 2.5-5 mm; fruiting pedicels 2.5-6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .19
18(17). Basal leaves pinnatisect with linear lobes; upper leaves linear to linear-oblanceolate; fruiting pedicels puberu-
lent adaxially; petals shorter than sepals or absent; style included in apical notch; Argentina . . . . . . . . . .L. spicatum
18. Basal leaves entire or serrate; upper leaves oblong; fruiting pedicels puberulent all around; petals subequaling
sepals; style subequaling apical notch; Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. cuzcoensis
19(17). Stems decumbent or ascending from base; rachis of raceme with spreading trichomes; Bolivia, Chile, Ecua-
dor, Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. chichicara
19. Stems erect at base; rachis of raceme with retrorse trichomes; Argentina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
20(19). Lower cauline leaves pinnatisect; petals 0.9-1.5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. parodii
20. Lower cauline leaves entire or dentate; petals 0.4-0.7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
21(20). Fruits obovate-suborbicular, 4.2-5.3 × 3.5-5 mm, apical notch 0.5-1 mm; basal leaves persistent, serrate-den-
tate; cauline leaves entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. hickenii
21. Fruits elliptic-ovate, 3.5-4.2 × 2.4-3 mm; apical notch 0.1-0.2 mm; basal leaves deciduous, pinnatifid; cauline lea-
ves dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. burkartii
22(1). Annuals or rarely biennials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23
22. Perennials with caudex, rarely subshrubs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .42
23(22). Fruits indehiscent; valves thick, prominently rugose to verrucose, closed and enclosing seeds . . . . . . . . . .24
23. Fruits dehiscent; valves thin, smooth or rarely reticulate, open . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25
24(23). Fruits reniform to ovate-cordate; valves prominently ridged; stamens 6; petals 1-2 mm . . . . . . .L. coronopus
24. Fruits didymous; valves without ridges; stamens 2; petals 0.4-0.5 mm . . . . . . . . . . . . . . . . . . . . . . . . .L. didymum
25(23). Fruit valves prominently reticulate-veined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26
25. Fruit valves smooth, not prominently veined28
26(25). Fruits 3.4-5 × 3.4-5 mm, winged all around . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. angustissimum
26. Fruits 2.2-3.3 × 2-3.2 mm, winged apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27
27(26). Sepals caducous; fruits subdidymous, usually wider than long, margin glabrous . . . . . . . .L. pseudodidymum
27. Sepals persistent; fruits not didymous, longer than wide; margin minutely puberulent, very
rarely glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. strictum
28(25). Stamens 6 or 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29
28. Stamens 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .31
29(28). Fruits 1.8-2 ×1.7-1.8 mm, apically wingless, valves pilose . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. pinnatifidum
29. Fruits 2.5-7 × 2-5.5 mm, apically winged; valves glabrous or very rarely puberulent along margin . . . . . . . . . .30
30(29). Fruiting pedicels strongly flattened, puberulent adaxially; stamens 4; cotyledons entire; style obsolete or very
rarely to 0.1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. nitidum
30. Fruiting pedicels terete or subterete, glabrous; stamens 6; cotyledons trifid; style 0.2-0.5(-0.8) mm . . .L. sativum
31(28). Fruits wingless; style exserted from apical notch of fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. cyclocarpum
31. Fruits apically winged; style included in apical notch of fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32
32(31). All leaves entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. horstii
32. At least some leaves dentate, incised, trifid, pinnatifid, or pinnatisect . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33
33(32). Fruits 1.5-2 × 1.3-1.5 mm, slightly reticulate; seeds neither winged nor margined . . . . . . . . . .L. myrianthum
33. Fruits 2-5 × 1.7-4.8 mm, not reticulate; seeds winged or margined, rarely neither . . . . . . . . . . . . . . . . . . . . . . .34
34(33). Middle or upper leaves or leaf segments linear, 0.5-2 mm wide; fruits 2-3 mm . . . . . . . . . . . . . . . . . . . . . .35
34. Leaves or leaf segments variously shaped, not linear, usually broader; fruits 2.5-5 mm . . . . . . . . . . . . . . . . . . .37
144
DARWINIANA 48(2) 141-167. 2010
35(34). Stems often decumbent; basal leaves 2-pinnatisect; fruits oblong-obovate, 1.7-2.1 mm wide;
Patagonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. filisegmentum
35. Stems erect; basal leaves 1-pinnatisect; fruits orbicular or orbicular-obovate, 2-3.2 mm wide; C and N. Argentina,
Paraguay . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .36
36(35). Fruiting pedicels terete, wingless, 2-3 mm; fruits 2.7-3.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. gracile
36. Fruiting pedicels flattened, narrowly winged, 2-6 mm; fruits 2-2.5 mm . . . . . . . . . . . . . . . . . . . .L. stuckertianum
37(34). Rachis of raceme papillate with straight subclavate trichomes . . . . . . . . . . . . . . . . . . . . . . . . . .L. densiflorum
37. Rachis or raceme puberulent or strigillose with recurved trichomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38
38(37). Fruits puberulent at least along margin when young . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. pubescens
38. Fruits almost always glabrous, if puberulent (L. argentinum) then leaves not pinnatisect . . . . . . . . . . . . . . . . . .39
39(38). Petals 1-2(-2.5) mm; fruits orbicular or suborbicular; cotyledons accumbent . . . . . . . . . . . . . . .L. virginicum
39. Petals absent or rudimentary and less than 1 mm; fruits obovate, obovate-oblong, elliptic, or rarely suborbicular;
cotyledons incumbent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40
40(39). Cauline leaves entire, dentate, laciniate, or rarely pinnatifid; fruits sharply acute winged apically, sparsely and
minutely puberulent on margin at least when young, or glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. argentinum
40. At least lower cauline leaves 1-3-pinnatisect or -pinnatifid; fruits obtusely winged apically, glabrous . . . . . . . .41
41(40). Fruits broadly elliptic-obovate to orbicular, (2.5-)3-3.5 mm, apical notch 0.1-0.3 mm;
widespread . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. bonariense
41. Fruits broadly obovate to oblong-obovate, (3-)3.7-4.5(-5) mm, apical notch 0.5-1 mm;
Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. costaricense
42(22). Subshrubs or perennial herbs with woody lower stems; basal leaves absent . . . . . . . . . . . . . . . . . . . . . . . . .43
42. Perennial herbs without woody stems; basal leaves present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .46
43(42). Fruits oblong or elliptic to obovate, not subdidymous, dehiscent, thin walled, smooth; style exserted from api-
cal notch; stamens 4; petals 1-3 mm44
43. Fruits orbicular to broadly obovate-suborbicular, subdidymous, indehiscent, breaking into closed, 1-seeded valves,
thick walled, reticulate-alveolate; style included in apical notch; stames 2 or 6; petals 0.3-0.6 mm . . . . . . . . . . . . .45
44(43). Petals white, 2.5-3 mm; fruits puberulent, 4-5 × 3-3.5 mm; Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . .L. beckii
44. Petals yellow, 1-1.5(-2) mm; fruits glabrous, 3-4 × 1.5-2.5 mm; Ecuador . . . . . . . . . . . . . . . . . . . . . . .L. quitense
45(43). Leaves entire or 3-5-lobed; stamens 2; fruits 2.5-3.5 × 3.5-4.5 mm, on terminal and lateral branches
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. rhytidocarpum
45. Leaves serrate; stamens 6; fruits 4-5 × 3-3.5 mm, axillary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. serratum
46(42). Plants scapose; cauline leaves absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .47
46. Plants not scapose; cauline leaves present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .48
47(46). Rachis and pedicels puberulent; stamens 6; fruits elliptic-rhombic, 4.5-6 mm, wingless, not notched;
Chile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. philippianum
47. Rachis and pedicels glabrous; stamens 4; fruits orbicular, 3.2-3.8 mm, winged apically, notched;
Bolivia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. solomonii
48(46). Stamens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .649
48. Stamens 2 or rarely . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .450
49(48). Plants 2-4 dm; fruits 5-7 × 3-4 mm; notch and style 0.25-0.5 mm, subequal; seeds 2-3 mm;
Brazil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. grandifructum
49. Plants (2-)3.5-12(-15) dm; fruits 1.6-2.7 × 1.3-1.8 mm, notch and style obsolete; seeds 0.8-1.3 mm; naturalized in
Argentina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. latifolium
50(48). Style exserted from or rarely subequaling apical notch of fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .51
50. Style distinctly included in apical notch of fruit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .57
51(50). Stamens 4; petals 1.2-2.2 mm wide; fruits reticulate, distinctly wider than long . . . . . . . . . . . . . . . .L. werffii
51. Stamens 2; petals 0.2-1.3 mm wide; fruits not reticulate, narrower than or rarely slightly wider than long . . . .52
52(51). Fruits 5-8 × 3-4 mm, 1.5-2 times as long as wide, puberulent at least when young, rarely glabrous; seeds blac-
kish 2-3 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. marginatum
52. Fruits 2.5-4.5(-5) × 2.5-5 mm, about as long as or slightly longer than or shorter than wide, glabrous; seeds brown,
1.5-2.2 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .53
53(52). Plants with appressed strongly crisped trichomes; stems often shorter than basal leavesL. jujuyanum
53. Plants with spreading to curved, non-crisped trichomes; stems longer than basal leaves . . . . . . . . . . . . . . . . . .54
54(53). Basal and lowermost cauline leaves serrate; fruits wingless, 4.5-5 mm wide, slightly wider than long
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. spathulatum
145
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
54. At least some basal and lowermost cauline leaves pinnatifid or pinnatisect; fruits narrowly winged apically, 2.5-
3.8(-4) mm wide, often longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .55
55(54). Petals (1-)1.5-2.5(-3) mm; style 0.3-1 mm; fruits 2.5-3.8(-4) mm wide, rhombic-suborbicular or rarely rhom-
bic-elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. meyenii
55. Petals 0.5-1.3 mm; style 0.1-0.3 mm; fruits 1.8-2.8 mm wide, elliptic or elliptic-obovate . . . . . . . . . . . . . . . . .56
56(55). Sepals persistent; fruits elliptic, 2.5-3.2 mm; stems 0.3-2.5 dm, decumbent; middle cauline leaves pinnatifid;
Argentina, Bolivia, Peru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. depressum
56. Sepals caducous; fruits elliptic-obovate, 3.8-4.5 mm; stems 3-5 dm, ascending; middle cauline leaves entire, sub-
apically toothed; Colombia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. trianae
57(50). Fruiting pedicels pubescent all around at least basally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .58
57. Fruiting pedicels glabrous or pubescent only adaxially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .60
58(57). Sepals caducous; fruits puberulent at least when young; petals 1.2-1.8 mm . . . . . . . . . . . . . .L. ecuadoriense
58. Sepals persistent or caducous; fruits glabrous; petals 0.6-1 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .59
59(58). Basal leaves pinnatifid with (2 or)3-8 lateral lobes on each side; middle cauline leaves serrate or laciniate;
fruits narrowly oblong-ovate to oblong, 2-2.3 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. abrotanifolium
59. Basal leaves entire or rarely with one lobe on each side; cauline leaves entire; fruits broadly ovate or obovate, 2.5-
3 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. fraseri
60(57). Petals spatulate to oblanceolate, 1.5-2 mm, longer than sepals; fruits 4-5.5 × 3-4 mm . . . . . .L. cumingianum
60. Petals linear, filiform or subulate, 0.2-0.8 mm, shorter than sepals; fruits 2-4(-4.2) × 1-3(-3.5) mm . . . . . . . . .61
61(60). Leaves or leaf lobes linear, entire, 0.3-1 mm wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .62
61. Leaves or leaf lobes pinnatifid, dentate, serrate, or incised, often broader . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .63
62(61). Fruiting racemes dense; fruiting pedicels puberulent adaxially; fruits orbicular to broadly elliptic, 2-2.5 mm
wide; C Argentina and Patagonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. spicatum
62. Fruiting racemes lax; fruiting pedicels glabrous; fruits elliptic, 1.8-2 mm wide; Bolivia . . . . . . . . .L. steinbachii
63(61). Fruits 2-2.8 × 1-1.8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .64
63. Fruits 3-4.2 × 2.4-4 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .65
64(63). Basal and lowermost cauline leaves serrate or dentate; stems 2-5 dm; racemes compact in fruit, rachis gla-
brous; fruiting pedicels recurved; Argentina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. boelckeanum
64. Basal and lowermost cauline leaves pinnatifid or pinnatisect; stems 0.3-1.5 dm; racemes elongated in fruit, rachis
pubescent; fruiting pedicels straight, appressed to rachis; Chile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. reichei
65(63). Stems single from base, erect; rachis of raceme with retrorse trichomes; seeds narrowly winged or margined;
Argentina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. burkartii
65. Stems few to seveal from base, decumbent to ascending; rachis of raceme with spreading trichomes; seeds win-
gless and marginless; elsewhere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .66
66(65). Stems 1.5-4.5 dm; fruits elliptic to obovate, glabrous; fruiting pedicels 2.5-5 mm; seeds 1.3-1.5 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. chichicara
66. Stems 0.4-1(-2) dm; fruits orbicular to ovate-orbicular, often puberulent along margin when young, fruiting pedi-
cels 1-2(-3) mm; seeds 1.5-1.9 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .L. rahmeri
Lepidium abrotanifolium Turcz., Bull. Soc.
Naturalistes Moscou 27: 308. 1854. TYPE:
Ecuador. Antisana, 14,000 ft, 1850, Jameson
s.n. (holotype KW; isotypes F!, G-BOIS!).
Lepidium abrotanifolium var. steinmannii Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 247. 1906.
TYPE: Bolivia. Oruro and Cochabamba, 4000 m,
1900, Steinmann s.n. (lectotype B! here designa-
ted).
Lepidium fraseri Thell. var. dielsianum O. E. Schulz ex
Diels, Biblioth. Bot. 116: 89. 1937. TYPE: Ecuador.
Chimborazo, N of Riobamba, Sanancajas, páramo,
3500 m, 28-VII-1933, L. Diels 400 (holotype B!).
Distribution. Bolivia, Ecuador, and Peru.
Observations. The lectotype designated herein is
the most complete of the two syntypes cited, and it
was named after its collector. The other syntype,
K. Fiebrig 2772 (B), was collected from Tarija,
Bolivia.
Lepidium angustissimum Phil., Anales Univ.
Chile 81: 333. 1892. TYPE: Chile. Travesía,
between Chañarcillo and Carrizal valley, IX-
1885, F. Philippi s.n. [lectotype SGO71482!
designated by M. Muñoz-Schick, Notic. Mens.
Mus. Nac. Hist. Nat. (Chile) 359: 7. 2007].
Lepidium tayloriae Al-Shehbaz, Novon 3: 93. 1993.
TYPE: Chile. Región III (Atacama), Copiapó Pro-
vince, between Huasco and Copiapó, ca. 20-25 km
W of Totoral on road to coaset, dry plains and hillsi-
de, 20°00' S, 70°50' W, 8-X-1991, C. M. Taylor, C.
von Bohlen & A. Marticorena 10804 (holotype MO!;
isotype CONC!).
Distribution. Endemic to Chile.
Observations. Upon the examination of the type
collections of Lepidium angustissimum and L. tay-
loriae, Muñoz-Schick (2007) concluded that they
are conspecific, a position with which I agree.
Lepidium appelianum Al-Shehbaz, Novon
12: 7. 2002. Hymenophysa pubescens C. A.
Mey. in Ledeb., Icon. Pl. 2: 20. 1830, non Lepi-
dium pubescens Desv., J. Bot. Agric. 3: 180.
1815, nec L. pubescens Tineo, Cat. Pl. Hort.
Panorm. 150. 1827. Cardaria pubescens (C. A.
Mey.) Jarm. in Keller et al., Weeds USSR 3: 29.
1934. TYPE: [Kazakhstan]. "Locis humidis
subsalsis deserti Soongoro-Kirghisici orientalis
versus montes Arkaul," 14-V-1826, C. A. Meyer
s.n. (lectotype LE designated by I. A. Al-Sheh-
baz et al., Novon 12: 7. 2002).
Distribution. Native to Central Asia, naturalized
in North America and South America (Argentina).
Lepidium argentinum Thell., Physis 9: 9.
1928. TYPE: Argentina. La Rioja, Chilecito,
Sañugasta, 30-I-1927, L. R. Parodi 7785 (lec-
totype BAA! here designated).
Lepidium bonariense L. var. stenocarpum Thell.,
Repert. Spec. Nov. Regni Veg. 11: 310. 1912. Lepi-
dium argentinum var. stenocarpum (Thell.) Thell.,
Physis 9: 10. 1928. TYPE: Argentina. La Rioja,
Entre la mina Jareta ya la altura del Espiritu Santo,
Sierra Famatina, 25-I-1879, G. Hieronymus & G. Nie-
derlein 785 (holotype B!; isotypes CORD![2], G-
DC!).
Lepidium argentinum var. virginicifolium Thell., Physis
9: 11. 1928. TYPE: Argentina. La Rioja, Chilecito,
Sañugasta, 30-I-1927, L. R. Parodi 7783 (lectotype Z
designated by C. L. Hitchcock, Lilloa 11: 105. 1945;
duplicates BAA!, GH!).
Distribution. Endemic to Argentina.
Observations. Thellung (1928) did not designate
a type for the species and recognized the two varie-
ties above. He listed under var. virginicifolium two
collections, Parodi 7785 and Parodi 7783, of
which Hitchcock (1945) took Parodi 7783 as the
varietal type. The only other collection examined
by Thellung is Parodi 7785, and the BAA sheet is
taken herein as the lectotype because it was not
possible to locate such collection among all of the
major European herbaria consulted.
Lepidium auriculatum Regel & Körn., Ind.
Sem. Hort. Petrop. 51. 1857. TYPE: Chile.
[Región XIV (Los Ríos). Prov. Valdivia], San
Juan, R. Philippi 317 (lectotype LE here desig-
nated; duplicates B!, G-BOIS!, P!, W!).
Lepidium calycinum Gordon, Mém. Acad. Montpel. 1:
416. 1853, syn. nov. Non Steph. ex Willd., Sp. Pl. 3:
433. 1800. Lepidium aletes J. F. Macbr., Candollea 5:
357. 1934. TYPE: France (adventive). Port-Juvenál
près Montpellier, 1853, Touchy s.n. (holotype
NCY!).
Lepidium araucanum Phil., Anales Univ. Chile 81: 335.
1892. Lepidium bipinnatifidum Desv. var. arauca-
num (Phil.) Reiche, Anales Univ. Chile 90: 95. 1895.
TYPE: Chile. Curanilahue, XI-1891, R. Philippi s.n.
(holotype SGO-071462!).
Lepidium pubescens Desv. var. fallax Thell., Bull. Herb.
Boissier ser. 2, 8: 913. 1908. TYPE: Chile. Quillota,
1829, C. G. L. Bertero 1080 (lectotype P! here desig-
nated).
Lepidium subvaginatum Steud. ex Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 249. 1906, syn.
nov. TYPE: Chile. Mt. Leona, Rancagua, 1828, C. G.
L. Bertero 364 (lectotype P! questionably designated
by C. L. Hitchcock, Lilloa 11: 112. 1945, and confir-
med herein; duplicates F!, G[2]!, GH[2]!, NY!, P!).
Distribution. Argentina, Bolivia, Chile, Para-
guay, and Uruguay.
Observations. The type collections of Regel's
new taxa are housed at LE, and the lectotype of
Lepidium auriculatum is not an exception. The
146
DARWINIANA 48(2) 141-167. 2010
sheet of L. pubescens var. fallax at P was the only
one annotated by Thellung as such, and it is desig-
nated herein as the lectotype of this taxon. Both
Thellung (1906) and Hitchcock (1945) recognized
Lepidium auriculatum, L. subvaginatum, and L.
calycinum (=L. aletes by Hitchcock) as distinct
species separated primarily by the degree of deve-
lopment of auricles on the cauline leaves, length of
nectar glands at the base of the fruit, and trichome
density on upper parts of the stem. However, the
study of substantial material reveals that a single,
vegetatively polymorphic species is involved. In
every floral, fruit, and seed morphology, the above
three species are basically indistinguishable. A
combination of annual habit, pinnatifid or pinnati-
sect and auriculate cauline leaves, persistent
sepals, and pedicels pubescent all around should
readily distinguish the species from the other
South American Lepidium.
The lectotype sheet of Lepidium pubescens var.
fallax includes five plants, of which three were
annotated by Thellung (1908) as this variety and
one each as L. pubescens and L. subvaginatum. A
close examination of all five plants reveals that
they belong to L. auriculatum.
Lepidium beckii Al-Shehbaz, Novon 9: 5. 1999.
TYPE: Bolivia. La Paz, Prov. José Romá de
Loayza, Baños Termales de Urmiri, 17°09' S,
68°05' W, 3500 m, 26-I-1996, S. G. Beck 21944
(holotype MO!; isotype LPB!).
Lepidium philippianum (Kuntze) Thell. var. boliviense
Thell., Neue Denkschr. Schweiz. Naturf. Ges. 41:
201. 1906, syn. nov. TYPE: Bolivia. La Paz, Chive-
sivi, Valeé de La Paz, 8500-12,500 ft, 1939, Pentland
s.n. (holotype P!).
Distribution. Endemic to Bolivia.
The type of var. boliviense is indistinguishable
from that of Lepidium beckii in every aspect of the
plant and clearly differs from L. philippianum in
which it was originally described (see couplet 42
of the key).
Lepidium bipinnatifidum Desv., J. Bot.
Agric. 3: 177. 1815. TYPE: Peru, sine data,
Dombey s.n. (holotype P!; isotype P!). The
holotype sheet was annotated in Desvaux's
handwriting.
Lepidium humboldtii DC., Syst. Nat. 2: 532. 1821.
TYPE: Ecuador, "Hab. locis aridis prope Chillo Qui-
tensium altit 1340 hexapod," A. J. A. Bonpland & F.
W. H. A. von Humboldt 2223 (holotype B!).
Senebiera dubia Kunth in Humboldt, Bonpland &
Kunth, Nov. Gen. Sp. 76. 1821 TYPE: Ecuador, sine
locus, F. W. H. A. von Humboldt & A. J. A. Bonpland
s.n. (holotype P; isotype B!).
Lepidium auritum Turcz., Bull. Soc. Imp. Naturalistes
Moscou 27: 307. 1854. TYPE: Ecuador. Quito,
Jameson 772 (holotype KW; isotype G-DEL!).
Lepidium sectifolium Steud., Flora 39: 412. 1856.
TYPE: Peru. Tabina, Lechler 1879 (holotype B!;
isotype K!).
Lepidium kalenbornii C. L. Hitchc., Lilloa 11: 88. 1945,
syn. nov. TYPE: Peru. Oroya near Luma, dry soil,
10,000-13,000 ft, A. S. Kalenborn 20 (holotype DS!;
isotypes GH!, MO!, NY!, US!).
Distribution. Bolivia, Colombia, Ecuador, Peru,
and Venezuela.
Observations. Hitchcock (1945) separated Lepi-
dium bipinnatifidum from L. kalenbornii by the
presence (vs. absence) of auricles, slightly longer
nectar glands (<0.3 vs. ca. 0.3 mm), and slightly
larger fruits (2.5-3.5 vs. 2-3 mm). However, these
differences are unrealistic, and he recognized both
auriculate and non-auriculate forms in L. bipinna-
tifidum and small (2-3 mm) and larger (3-4 mm)
fruits in L. kalenbornii (see key couplets 6 and 47,
respectively).
Lepidium boelckeanum A. Prina, Hickenia
2(18): 81. 1993. TYPE: Argentina. La Pampa,
Depto. Chapaleufú, entre B. Larroudé y Río V,
29/30-X-1984, H. O. Troiani & A. O. Prina
8193 (holotype SRFA!; isotype BACP!).
Distribution. Endemic to Argentina.
Lepidium bonariense L., Sp. Pl. 2: 645. 1753.
TYPE: "Habitat in Bonaria" (lectotype designa-
ted by W. Marais, Flora of Southern Africa 13:
93. 1970: "Thlaspi Bonar. multiscissum, flore
147
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
148
DARWINIANA 48(2) 141-167. 2010
invisibili" in Dillenius, Hort. Eltham., 2: 381, t.
286, f. 370. 1732).
Thlaspi multifidum Poir., Encycl. 7: 545. 1806. TYPE:
Uruguay. Montevideo, 1767, P. Commerson s.n.
(holotype P; isotypes P[3]!). The sheet annotated by
Poiret is taken herein as the holotype.
Lepidium mendocinum Phil., Anales Univ. Chile 36:
160. 1870. TYPE: Argentina. Inter Mendocino et
Santiago, P. Ortega & E. Reed s.n. (holotype SGO-
045144).
Lepidium bonariense var. hirsutulum Thell., Repert.
Spec. Nov. Regni Veg. 11: 310. 1912. TYPE: Argen-
tina. Córdoba, without locality, 1876, G. Hieronymus
& G. Niederlein s.n. (holotype B!).
Lepidium bonariense var. microcarpum Thell., Repert.
Sp. Nov. Regni Veg. 13: 302. 1914. TYPE: Argenti-
na. Córdoba, Depto. Capital: prope ciudad de Córdo-
ba, T. Stuckert 5647 (holotype G; isotype CORD!).
Lepidium bonariense var. pseudovirginicum Thell.,
Repert. Sp. Nov. Regni Veg. 13: 301. 1914. TYPE:
Argentina. Córdoba, Punilla, Los Cocos, T. Stuckert
19614 (lectotype G-DEL! here designated; duplicate
CORD!).
Distribution. Argentina, Bolivia, Brazil, Chile,
Paraquay, and Uruguay; naturalized in Australia,
Europe, and South Africa.
Observations. Stuckert 19614 at G was desig-
nated herein as the lectotype because it is the
most complete specimen that agrees with the des-
cription and because it was annotated as that
name by its author. Thellung (1906, 1914) and
Hitchcock (1945) broadly delimited Lepidium
bonariense to include at least three additional
species, including L. gracile, L. stuckertianum,
and L. tandilense. These species are easily distin-
guished by the key above.
Lepidium burkartii Boelcke, Parodiana 3(1):
21. 1984. TYPE: Argentina. Entre Ríos, Depto.
Gualeguaychú, sotobosque del río Gualeguay-
chú, pocos km al N de Gualeguaychú, 5-XI-
1983, N. S. Troncoso, N. Bacigalupo & N. Tur
3740 (holotype SI!; isotypes BACP!, SI!). The
holotype and isotype sheets at SI are marked a
and b, respectively. The holotype sheet has two
plants and the isotype has four.
Distribution. Endemic to Argentina.
Lepidium campestre (L.) W. T. Aiton, Hortus
Kew., ed. 2, 4: 88. 1812. Thlaspi campestre L.,
Sp. Pl. 2: 646. 1753. TYPE: "Habitat in Euro-
pae arvis, viis argillosis, apricis" [lectotype
Herb. Linn. 825.8 (LINN) here designated].
Distribution. Native to Europe, introduced into
North America, South America (Chile), Australia,
and South Africa.
Observations. Jarvis (2007) did not designated a
lectotype among the five original material he listed
under Thlaspi campestre. The specimen placed on
Linnaean website (http://www.linnean-
online.org/view/plants_alpha/thlaspi_campestre.ht
ml) is designated herein as the lectotype.
Lepidium chalepense L., Cent. Pl. II, 23. 1756.
Cardaria chalepensis (L.) Hand.-Mazz., Ann.
Nat. Hofmus. Wien 27: 55. 1913. Cardaria
draba subsp. chalepensis (L.) O. E. Schulz in
Engler & Prantl, Nat. Pflanzenfam., ed. 2, 17b:
417. 1936. TYPE: "Habitat in Oriente" [lectoty-
pe Herb. Linn. No. 824.20 (LINN) designated
by S. M. H. Jafri, in Nasir & Ali, Fl. W. Pakis-
tan 55: 68. 1973].
Distribution. Native to SW Asia; naturalized in
Europe and North and South America (Argentina).
Lepidium chichicara Desv., J. Bot. Agric. 3:
179. 1815. TYPE: Peru. Chichicara, Dombey
s.n. [as Dombrey] (holotype P!; isotype P!). The
sheet annotated by Desvaux is taken herein as
the holotype.
Lepidium lanceolatum Walp., Nov. Actorum. Acad.
Caes. Leop.-Carol. Nat. Cur. 19 (Suppl. 1): 250.
1843, non Presl, Fl. Sicul. 1: 82. 1826. Lepidium chi-
chicara var. lanceolatum (Walp.) Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 221. 1906.
Lepidium walpersii J. F. Macbr., Candollea 5: 357.
1934. TYPE: Peru. Lago Titicaca, 12400 ft, F. J . F.
Meyen s.n. (holotype B!).
Lepidium chichicara var. pseudobipinnatifidum Thell.,
Neue Denkschr. Schweiz. Naturf. Ges. 41: 221.
1906. TYPE: Ecuador, sine locus, 1871, A. Sodiro 58
(holotype B!).
Lepidium chichicara var. rhombocarpum Thell., Repert.
149
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
Spec. Nov. Regni Veg. 11: 309. 1912. TYPE: Boli-
via. La Paz, Palca, Illimani, 3600-4800 m, 1906, R.
Hauthal 268 (holotype B!).
Distribution. Bolivia, Chile, Ecuador, and Peru.
Lepidium coronopus (L.) Al-Shehbaz, Novon
14: 156. 2004. Cochlearia coronopus L., Sp. Pl.
2: 648. 1753. TYPE: "Habitat in Europae apri-
cis, nudis" [lectotype Herb. Linn. No. 826.5
(LINN) designated by B. Jonsell & C. Jarvis,
Nord. J. Bot. 22: 68. 2002].
Lepidium squamatum Forssk., Fl. Aegypt. Arab. 117.
1775. Coronopus squamatus (Forssk.) Aschers., Fl.
Prov. Brandenb. 62. 1860. TYPE: [Egypt], Alexan-
dria, P. Forsskål s.n. (holotype C).
Distribution. Native to Eurasia and N Africa;
naturalized in North America, South America
(Chile), Australia, and South Africa.
Lepidium costaricense Thell., Bull. Herb.
Boissier, ser. 2, 4: 713. 1904. TYPE: Costa
Rica. San José, XI-1875, H. Polakowsky 533
(lectotype Z! here designated; duplicate W!).
Lepidium costaricense var. friedrichsthalii Thell., Neue
Denkschr. Schweiz. Naturuf. Ges. 41: 252. 1906.
TYPE: Guatemala. Jinotepe, 1841, Friedrichsthal
1199 (holotype W).
Lepidium virginicum L. subsp. centrali-americanum
Thell., Neue Denkschr. Schweiz. Naturf. Ges. 41:
225. 1906. Lepidium virginicum var. centrali-ame-
ricanum (Thell.) C. L. Hitchc., Madroño 8: 128.
1945. TYPE: Mexico. Yucatán, Izamal, 1895, G. F.
Gaumer 456 (lectotype B! here designated; dupli-
cate G!).
Distribution. Belize, Colombia, Costa Rica, El
Salvador, Guatemala, Honduras, Mexico, Nicara-
gua, and Panama.
Observations. The reasons why the lectotypes of
Lepidium costaricense and L. virginicum subsp.
centrali-americanum were designated herein are
because they were annotated by Thellung and
were among the most complete specimens exami-
ned by the present author.
Lepidium crassius (C. L. Hitchc.) Al-Shehbaz,
comb. nov. Lepidium cyclocarpum var. crassius
C. L. Hitchc., Lilloa 11: 94. 1945. TYPE: Peru.
Arequipa, 40 Km S of Chala, 5 km from the
sea, 700 m, 22-IX-1938, C. R. Worth & J. L.
Morrison 15701 (holotype MO!; isotypes G!,
GH, MO[2]!, NA!, UC!).
Distribution. Endemic to Peru.
Observations. Although Hitchcock (1945) des-
cribed this species as a variety of Lepidium cyclo-
carpum, the differences between them are so
substantial that it is difficult to imagine why there
were treated as conspecific. Lepidium crassius
differs from L. cyclocarpum by being a densely
pubescent perennial (vs. sparsely pubescent
annual) with hirsute (vs. puberulent) stems 3-5
(vs. 0.5-2) dm tall, auriculate (vs. non auriculate)
middle and upper cauline leaves, narrowly win-
ged fruiting pedicels pubescent all around (vs.
wingless pedicels puberulent only adaxially),
obovate petals , 2-3 × 1-1.8 mm (vs. spatulate to
oblanceolate petals 1-1.5 × 0.2-0.5 mm), four (2
median and 2 lateral) stamens (vs. only 2 median),
nectar glands 0.5-0.7 mm (vs. 0.1-0.2 mm) long,
fruits without (vs. with) apical notch, and styles
0.7-0.1 (vs. 0.2-0.4) mm.
Lepidium cumingianum Fisch. & C. A. Mey.,
Index Sem. Hort. Petrop. 1: 30. 1835. TYPE:
Chile, sine locus, 1834, H. Cuming s.n. (holoty-
pe LE).
Lepidium cumingianum subsp. berteroanum Thell.,
Neue Denkschr. Schweiz. Naturf. Ges. 41: 218.
1906. TYPE: Chile. [Región V]: Valparaíso, 1829, C.
G. L. Bertero 1082 (lectotype G-DC! here designa-
ted; duplicates F!, GH!, NY!, P!).
Lepidium cumingianum subsp. orbiculatum Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 219. 1906.
TYPE: Chile. [Región I], Tarapacá, R. Philippi s.n.
(lectotype B! here designated; duplicate W!).
Lepidium cumingianum var. canescens Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 219. 1906.
TYPE: Chile. Santiago, 1818, C. G. L. Bertero 366
(holotype G-DEL!).
Lepidium cumingianum var. subsagittatum Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 219. 1906.
TYPE: no specimens cited.
Distribution. Endemic to Chile.
Observations. The designated lectotypes of
Lepidium cumingianum subsp. orbiculatum and
subsp. berteroanum are represented by more
duplicates than the others cited by Thellung, and
all were examined by the present author. Hitch-
cock (1945) indicated that Bertero 367 is the type
collection of Lepidium cumingianum, but there is
no indication that supports that. No collection data
were provided for var. subsagittatum, and only
"Herb. Petrop," now LE, was given.
Lepidium cuzcoensis Al-Shehbaz, sp. nov.
TYPE: Peru. Depto. Cuzco, Prov. Urobamba,
Laderas de Muyock, 2860 m, 18-I-1963, C.
Vargas 14111 (holotype US; isotype US). Fig.
1.
Species perennis, staminis 2, foliis basalibus
integris vel serratis, petalis albis obovato-orbicu-
laris 0.7-1 mm longis et sepalis equilongis, fructi-
bus ellipticis 2.5-3 mm longis, apice emarginatis
stylis equilongibus a congeneribus diversa.
Herbs, perennial, puberulent with straight, spre-
ading trichomes 0.01-0.1 mm; caudex simple or
few branched, slender. Stems 4-13 cm, decumbent
to ascending, few from caudex, branched above.
Basal leaves 1.5-3 cm; petioles 0.6-1.5 cm, persis-
tent, only slightly flattened at base; blade oblance-
olate, 2-5 mm wide, entire or serrulate, puberulent
at margin; lowermost cauline leaves shortly petio-
late to subsessile, not auriculate at base; middle
and upper cauline leaves entire or few toothed,
oblong, minutely auriculate at base. Racemes
dense, slightly or considerably elongated in fruit;
rachis densely and minutely puberulent with
straight trichomes; fruiting pedicels ascending at
base, recurved, divaricate, 2-3 mm, narrowly win-
ged, minutely puberulent all around. Sepals bro-
adly ovate, 0.8-1.1 mm, somewhat persistent, with
a broadly white margin and apex, sparsely puberu-
lent; petals white, broadly obovate-suborbicular,
0.7-1 × 0.6-0.9 mm; stamens 2, median; filaments
0.6-1 mm; anthers ca. 0.2 mm. Fruits dehiscent,
broadly elliptic, 2.5-3 × 2.2-2.5 mm, glabrous, not
veined, narrowly winged apically, apex emargina-
te; apical notch 0.15-0.2 mm; style 0.15-2 mm,
equaling apical notch. Seeds brown, ovate-oblong,
wingless, 1.1-1.2 × 0.7-0.8 mm; cotyledons
incumbent.
Etymology. The species is named for the Peru-
vian province of Cuzco.
Distribution. Known only from the type collec-
tion.
Observations. Lepidium cuzcoensis does not
appear to be closely related to any of the South
American species. It is readily distinguished by a
combination of perennial habit, decumbent or
ascending stems, minutely puberulent pedicels all
around, two stamens, broadly obovate-orbicular
petals 0.7-1 mm subequaling sepals, and broadly
elliptic fruits 2.5-3 mm long with short style equa-
ling the apical notch.
Lepidium cyclocarpum Thell., Neue Denkschr.
Schweiz. Naturf. Ges. 41: 214. 1906. TYPE:
Peru. Lima, Amancaes Hills, Matthews (as
Mathew) 752 (lectotype W! designated by J. F.
Macbride, Publ. Field Mus. Nat. Hist., Bot ser.
13(2): 949. 1938; isolectotype K!).
Distribution. Endemic to Peru.
Observations. Hitchcock (1945) indicated that
the type collection is Weberbauer 1614 and over-
looked the earlier lectoypification given above.
Lepidium densiflorum Schrad., Index Sem.
Hort. Gotting. 4. 1832. TYPE: Germany. Sine
locus, 1831, Schrader s.n. (holotype Z!).
Lepidium neglectum Thell., Bull. Herb. Boissier, ser. 2,
4: 708. 1904. TYPE: United States of America. New
York, New York City, Bedford Park, 12-VIII-1899,
Wilson s.n. (lectotype Z! here designated).
Distribution. Native to North America, introdu-
ced into South America (Argentina), and Europe.
Observations. The above lectotype of Lepidium
neglectum is the only sheet of the species annota-
ted by Thellung and housed in the institution
150
DARWINIANA 48(2) 141-167. 2010
where he worked. Lepidium densiflorum was first
reported from Argentina by Prina (1997).
Lepidium depressum Thell., Neue Denkschr.
Schweiz. Naturf. Ges. 41: 201. 1906. TYPE:
Bolivia. [La Paz], Larecaja, Omasuyos, vicinity
of Achacache, Cerro Avichaca, 4000-4200 m,
Jan-May 1861, G. Mandon 922 (lectotype G!
here designated; duplicate W!).
Distribution. Argentina, Bolivia, and Peru.
Observations. The reason why the above sheet
of Lepidium depressum is designated herein as the
lectotype is because it is the most complete of
those annotated by Thellung.
Thellung (1906) gave the collection year for the
type collection as 1857, but the lectotype has the
year as 1861, and none of that original collection
carry the date as he stated.
Lepidium depressum is reported herein for the
first time from Argentina and Peru. All of the
Argentinean records above were previously unde-
termined, whereas the Peruvian record was cited
by Hitchcock (1945) as L. weddellii, a name that
he mishandled and treated herein as a synonym of
L. meyenii.
Specimens examined
ARGENTINA. Jujuy. Depto. Humahuaca,
Mina Aguilar, Cabrera et al. 18949 (LP). Depto.
Yavi, La Quiaca, Cabrera et al. 15294 (LP). La
Rioja. Depto. Famatina, Sierra de Famatina,
camino a la mina La Mexicana, Kiesling et al.
6290 (BAA). Tucumán. Sierra Aconquija,
Cerro Muñoz, Lillo 4163 (GH, LIL, US). Depto.
Tafí, Co. El Negrito, Giusti et al. MC545
(BAA).
PERU. Puno. Chuquibambilla, Pennell 13396
(F, GH, NY).
Lepidium didymum L., Syst. Nat., ed. 12, 2:
433. 1767. Coronopus didymus (L.) Sm., Fl.
Brit. 2: 691. 1804. Senebiera didyma (L.) Pers.,
Synop. Pl. 2(1): 185. 1806. TYPE: [lectotype
Herb. Linn. No. 824.16 (LINN!) designated by
W. Fawcett & A. B. Rendle, Flora of Jamaica 3:
244. 1914].
Senebiera pinnatifida DC., Mem. Soc. Hist. Nat. Paris
1799: 144, t. 9. 1799. TYPE: not designated.
Coronopus didymus var. macrocarpus Muschl., Bot.
Jahrb. Syst. 41: 137. 1908. TYPE: Uruguay. Concep-
ción, P. G. Lorentz 236 (lectotype B! here designa-
ted).
Coronopus didymus var. procumbens Muschl., Bot.
Jahrb. Syst. 41: 137. 1908. TYPE: Argentina, Ciéne-
ga, 10/17-I-1874, P. G. Lorentz & G. Hieronymus 650
(lectotype B! here designated).
Coronopus leptocarpus Boelcke, Darwiniana 19: 395.
1975, syn. nov. TYPE: Chile. Concepción, Depto.
Lautaro, Laraquete, 11-XI-1955, O. Boelcke 7309
(holotype SI!; isotype BAA!).
Coronopus leptocarpus var. microcarpus Boelcke, Dar-
winiana 19: 397. 1975, syn. nov. TYPE: Chile. Curi-
có, Depto. Vichuquén, Llico, 20-X-1938, E. Barros
2722 (holotype SI!).
Distribution. Native to southern South America,
naturalized in Central and North America, Europe,
Asia, Africa, and Australia.
Observations. The lectotypes of Coronopus
didymus var. procumbens and var. macrocarpus
were designated as such because they were the
only sheets examined by the present author and
annotated by Muschler among all the collections at
B, where he deposited his types. None of the ori-
ginal material cited under Senebiera pinnatifida
was examined by the present author and, therefo-
re, the species remains untypified.
Lepidium draba L., Sp. Pl. 2: 645. 1753. Carda-
ria draba (L.) Desv., J. Bot. Agric. 3: 163.
1815. TYPE: "Habitat in Germania, praesertim
Austria, Gallia, Italia", "Herb. Clifford: 331.
Lepidium 2, sheet 2" (lectotype BM-000646273
designated by B. Jonsell & C. Jarvis, Nord. J.
Bot. 22: 70. 2002).
Distribution. Native to Eurasia; naturalized in
South Africa, Australia, North America, and South
America (Argentina, Bolivia, Chile, and Uru-
guay).
151
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
Lepidium ecuadoriense Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 222. 1906.
TYPE: Ecuador. Cotopaxi, páramo, 4100 m,
VII-1903, Hans Meyer 198 (holotype JE!).
Distribution. Endemic to Ecuador.
Observations. Hitchcock (1945) reduced Lepi-
dium ecuadoriense to synonymy of L. chichicara,
but the two are so different in habit, indumentum,
and fruits that justify their recognition as distinct
species. From the latter, L. ecuadoriense differs by
having thick (vs. thin) caudex, shorter stems (0.5-
1.5 vs. 1.5-4.5 dm), non-auriculate (vs. often auri-
culate) cauline leaves, fruiting pedicels puberulent
all around (vs. only adaxially), petals subequaling
or slightly longer (vs. shorter) than sepals, puberu-
lent (vs. glabrous) fruits, and styles subequaling
(vs. distinctly shorter than) apical notch.
Lepidium filisegmentum C. L. Hitchc., Lilloa
11: 125. 1945. TYPE: Argentina. Santa Cruz,
Caleta Olivia, 100-200 m, 12-X-1929, A. Donat
185 (holotype GH!; isotypes BAA!, CAS!, F!,
GH!, MO!, NY!, SI!, UC!).
Lepidium reticulatum Howell var. austro-americanum
Thell., Bull. Herb. Boiss. 1908: 814. 1908. TYPE:
Argentina. Región entre lago Buenos Aires Norte y
Codo Río Mayer, 46º10' -48º15' S, 71-72º20' W, Río
Lista, 700 m, 5-III-1903, L. Platen & U. Greiner 60
(holotype Z; isotype SI!).
Distribution. Patagonian Argentina and Chile.
Lepidium fraseri Thell., Neue Denkschr.
Schweiz. Naturf. Ges. 41: 217. 1906. Lepidium
abrotanifolium Turcz. var. fraseri (Thell.) C. L.
Hitchc., Lilloa 11: 91. 1945. TYPE: Ecuador.
Sine locus, 1860, Fraser s.n. (holotype G-DC!).
Lepidium fraseri subsp. decipiens Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 217. 1906.
TYPE: Ecuador. Pichincha, 3000-3800 m, 1873, A.
Sodiro 55 (holotype B!).
Distribution. Endemic to Ecuador.
Observations. Lepidium fraseri is a very distinc-
tive species reduced by Hitchcock (1945) to a
variety of L. abrotanifolium. From the latter, it dif-
fers by always having glabrous (vs. puberulent),
entire (vs. pinnatifid) basal leaves 0.5-3 mm (vs. 7-
15 mm wide), entire (vs. 3-5-lobed) cauline lea-
ves, and broadly ovate to obovate (vs. oblong to
narrowly oblong-ovate) fruits 2.5-3 (vs. 2-2.3) mm
wide.
Lepidium gracile (Chodat & Hassl.) Boelcke,
Parodiana 4(1): 36. 1986. Lepidium bonariense
L. var. gracile Chodat & Hassl., Bull. Herb.
Boissier 2, 3: 795. 1903. Lepidium calycinum
var. gracile (Chodat & Hassl.) Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 245. 1906.
TYPE: Paraguay. Concepción, 1901/1902, E.
Hassler 7545 (holotype G!; isotypes G!, GH!,
P!, UC!).
Distribution. Argentina and Paraguay.
Observations. Thellung (1906) treated this spe-
cies as a variety of Lepidium bonariense, whereas
Hitchcock (1945) reduced it to synonymy of the
latter. As shown by Boelcke (1986) the differences
between the two species are substantial to merit
their recognition as independent.
Lepidium grandifructum C. L. Hitchc., Lilloa
11: 82. 1945. TYPE: Brazil, Upper Río Negro
River, 3-III-1906, Weiss & Schmidt s.n. (holoty-
pe NY!).
Distribution. Endemic to Brazil and known thus
far only from the type collection.
Lepidium heterophyllum Benth., Cat. Pl. Pyré-
nés 95. 1826. TYPE: France. Valleé d'Eynes,
sine data, no collector name or date (holotype
K!).
Distribution. Native to Europe; naturalized in
North America and South America (Chile).
Lepidium hickenii Al-Shehbaz, sp. nov. TYPE:
Argentina. Buenos Aires, Mar del Plata, puerto,
152
DARWINIANA 48(2) 141-167. 2010
153
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
Fig. 1. Lepidium cuzcoensis. A, plant. B, sepal. C, petal. D, stamen. E, fruit. Scales: A = 1 cm, B-E = 1 mm. Drawn
by the author from C. Vargas 14111 (holotype US).
cantera & altiplano, 15-XI-1932, C. M. Hicken
s.n. (holotype SI-21077b; isotypes SI-21077a,
c). Fig. 2.
Differt a Lepidio burkartii et L. parodii foliis
basalibus serrato-crenatis, fructibus obovato-sub-
orbicularis 3,5-5 mm latis, reticulatis, apice emar-
ginatibus 0,5-1 mm longis.
Herbs, perennial, with thick, simple or few-
branched caudex. Stems 10-45 cm, erect, striate,
often simple, glabrous below, strigillose above,
simple or divaricately branched above. Basal lea-
ves with petioles 1-2 cm, persistent well after fruit
maturity; leaf blade elliptic-lanceolate, 2-4 × 1-2
cm, densely strigillose, serrate-crenate, not ciliate;
cauline leaves sessile, 1-1.8 × 0.2-0.4 cm, linear-
lanceolate, auriculate, entire, densely strigillose,
uppermost sparsely so. Racemes dense in fruit;
rachis retrorsely strigillose; fruiting pedicels
slightly arcuate to nearly straight, divaricate-
ascending, 4-6 mm, strigillose adaxially. Sepals
ovate, 0.8-1 mm, margin white, subapically strigi-
llose, caducous; petals white, linear, 0.5-0.7 mm;
stamens 2, median; filaments ca. 0.5 mm; anthers
0.2-0.3 mm. Fruits dehiscent, obovate-suborbicu-
lar, 4.2-5.3 × 3.5-5 mm, glabrous, slightly reticula-
te, winged apically, apex deeply emarginate; api-
cal notch 0.5-1 mm; style obsolete, included in
apical notch. Seeds brown, ovate, narrowly win-
ged or margined, ca. 2 × 1.5 mm; cotyledons
incumbent.
Etymology. Lepidium hickenii is named in honor
of Cristóbal María Hicken (1 January 1875 - 11
March 1933), a renowned Argentinean botanist
who collected the type cited above.
Distribution. Nothing is known about the habitat
in which the species grows. It appears to be highly
restricted in Buenos Aires Province.
Observations. From the closely related perennial
and Argentinean-endemic Lepidium with auricula-
te cauline leaves, L. hickenii is easily distinguished
from L. burkartii and L. parodii by having serrate-
crenate (vs. pinnatifid or pinnatisect) basal leaves,
broader, obovate-orbicular fruits 3.5-5 mm wide
(vs. elliptic-ovate to obovate fruits 2.4-3.5 mm)
with, reticulate (vs. smooth) valves, and deeper
apical notch 0.5-1 (vs. 0.1-0.3) mm. The new spe-
cies also resembles L. pedersenii in having undivi-
ded basal and cauline leaves. However, it is readily
distinguished by its perennial (vs. annual) habit,
densely strigillose leaf surfaces (vs. glabrous or
puberulent midvein and leaf margin), longer frui-
ting pedicels 4-6 (vs. 2-3) mm, caducous (vs. per-
sistent) sepals, larger fruits 4.2-5.3 × 3.5-5 mm
(vs. 2.5-3.5 × 2-3 mm), and deeper apical fruit
notch 0.5-1 (vs. 0.4-0.5) mm.
Paratypes
ARGENTINA. Buenos Aires. Partido Neco-
chea, 19-XII-1944, Rodríguez 857 (LIL); Partido
Guaminí, Guaminí, Laguna del Monte, Nicora
4214 (SI); Partido Gral. Pueyrredón, Mar del
Plata, Hicken 2304 (SI).
Lepidium horstii Johow ex Skottsb., Handl. K.
Vetensk. Vitterh.-Samhales., Goteborgs V. ser.
B, 6: 33. 1937. TYPE. Chile. [Región V (Valpa-
raíso)]: San Ambrosio Island, 7 X 1896, F. R. A.
Johow s.n. (holotype not seen).
Distribution. Endemic to Chile.
Lepidium johnstonii C. L. Hitchc., Lilloa 11:
108. 1945. TYPE: Chile. [Región II (Antofa-
gasta)], Prov. Antofagasta, 5-6 km NE of Taltal,
300 m, 14-X-1938, C. R. Worth & J. L. Morri-
son 15845 (holotype UC!; isotype DS!).
Distribution. Endemic to Chile.
Lepidium jujuyanum Al-Shehbaz, Ann. Mis-
souri Bot. Gard. 76: 1189. 1989. TYPE: Argen-
tina. Jujuy, Depto. Humahuaca, Tres Cruces,
3750 m, 6-IV-1973, Barbara Ruthsatz 506/7
(holotype GH!; isotype BAA!).
Distribution. Endemic to Argentina.
Lepidium latifolium L., Sp. Pl. 2: 644. 1753.
TYPE: "Habitat in Galliae, Angliae umbrosis,
154
DARWINIANA 48(2) 141-167. 2010
succulentis" [lectotype Herb. Linn. No. 824.11a
(LINN) designated by S. M. H. Jafri, in Nasir &
Ali, Fl. W. Pakistan 55: 60. 1973].
Distribution. Native to Africa, Asia, and Europe;
naturalized in North America, South America
(Argentina), Australia.
Lepidium marginatum Griseb., Abh. König. Ges.
Wiss. Göttingen. 19: 72. 1874. Lepidium meyenii
subsp. marginatum (Griseb.) Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 204. 1906.
TYPE: Argentina. Catamarca, Sierra de Belen,
Vayas altas, 9-11,000 ft, I-1872, P. G. Lorentz 598
(holotype GOET!; isotypes B!, CORD![2]).
Lepidium boelckei Al-Shehbaz, Ann. Missouri Bot.
Gard. 76: 1189. 1989, syn. nov. TYPE: Argentina.
Jujuy, Depto. Humahuaca, Cerro La Soledad, 3500
m, 23-I-1929, S. Venturi 8859 (holotype US!; isotype
LIL!).
Distribution. Endemic to Argentina.
Observations. Hitchcock (1945) treated Lepi-
dium marginatum as a synonym of L. meyenii, but
the species is substantially distinct for laving lan-
ceolate to elliptic and puberulent (vs. rhombic to
rhombic-elliptic or rhombic-suborbicular and gla-
brous) fruits 5-8 [vs. (2.5-)3-4.5(-5)] mm, fruiting
pedicels puberulent all around (vs. often only ada-
xially), strongly curved (vs. usually straight) tri-
chomes, and at least some undivided (vs. always
pinnatifid or pinnatisect) basal leaves. Thellung
(1906) reduced L. marginatum to a subspecies of
L. meyenii, but the differences above strongly sup-
port the recognition of both as distinct species.
When Lepidium boelckei was described over 20
years ago (Al-Shehbaz, 1989), the type collection
and adequate material of L. marginatum were
available for that study. Both Hitchcock (1945)
and Thellung (1906) did not recognize the latter as
a distinct species. The study herein of ample
collections clearly reveals that the two plants are
conspecific.
Lepidium meyenii Walp., Nov. Act. Nat. Cur.
19. Suppl. 1: 249. 1843. TYPE: Peru. "In planis
circa Pisacomam," 15,000 ft, IV-1831, F. J . F.
Meyen 33 (holotype B!).
Lepidium gelidum Wedd., Ann. Sci. Nat. V, 1: 283.
1864. Lepidium meyenii subsp. gelidum (Wedd.)
Thell., Neue Denkschr. Schweiz. Naturf. Ges. 41:
203. 1906. Lepidium meyenii var. gelidum (Wedd.)
Hosseus, Bol. Acad. Nac. Ci. Córdoba 26: 101. 1921.
TYPE: Bolivia. Chuquisaca, Prov. Cinti, I-1846, H.
A. Weddell 3955 (lectotype P! designated by A. The-
llung, Neue Denkschr. Schwiz. Natürf. Ges. 41: 203.
1906; duplicate P!).
Lepidium affine Wedd., Ann. Sci. Nat., Bot. sér. 5, 1:
284. 1864, non Lepidium affine Ledeb., Index Sem.
Hort. Dorpat. 22. 1821. Lepidium meyenii Walp. var.
affine Thell., Neue Denkschr. Schweiz. Naturf. Ges.
41: 204. 1906. Lepidium weddellii O. E. Schulz,
Notizbl. Bot. Gart. Berlin-Dahlem 11: 391. 1932.
TYPE: Bolivia. Prov. Omasuyos, vicinis Achacace,
Taypichuru, in glareosis, 4000 m, 1857, G. Mandon
927 (lectotype P! partially designated by C. L. Hitch-
cock, Lilloa 11: 86. 1945, and herein; duplicates G-
BOIS!, G-DC!).
Lepidium peruvianum G. Chacón, Revista Peru. Biol.
3(2): 202. 1990, syn. nov. TYPE: Peru. Pasco, Depto.
Pasco, Huarancaca, dentro de la ciudad de Cerro de
Pasco, 4300 m, 9-IX-1989, G. Chacón s.n. (holotype
USM; isotypes B!, UC!, Z!).
Distribution. Argentina, Bolivia, and Peru.
Observations. The cultivated Lepidium of the
Peruvian highlands, better known as "maca," have
correctly been assigned to L. meyenii by various
authors. However, their recognition by Chacón
(1990) as L. peruvianum has no morphological
support other than the production of fleshy roots in
the cultivated (vs. nonfleshy roots in the wild)
forms of this species. One can draw so many simi-
lar parallels among the cultivated vs. wild forms of
radish (Raphanus sativus L.), turnip (Brassica rapa
L.), and rape (B. napus L.), all of which produce
fleshy roots in cultivation but do not produce them
when they become naturalized. Therefore, L. peru-
vianum does not have any merits. L. weddellii J. F.
Macbr., Candollea 5: 357. 1934, is nom. superfl.
Lepidium myrianthum Phil., Anales Mus. Nac.
Chile 8: 5. 1891. Lepidium ruderale L. var.
myrianthum (Phil.) Reiche, Anales Mus. Nac.
Chile 90: 96. 1896. TYPE: Chile. Antofagasta,
155
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
Cueva de Colorado, 23-I-1885, F. Philippi 1825
(lectotype SGO-64001 here designated).
Lepidium spicatum f. microcarpum Hicken, Physis
(Buenos Aires) 2: 13. 1915. TYPE. Argentina. Río
Negro, vicinity of General Roca, 250-360 m, IX-
1914 / II-1915, W. Fischer 37 (holotype SI!; isotypes
F[2]!, GH[2]!, K!, MO[2]!, NY!, US!).
Distribution. Argentina, Chile.
Observations. The original description of Lepi-
dium myrianthum was based on collections made
at 3880 m from Cueva de Colorado. Only two Phi-
lippi sheets from that locality are at SGO, and the
label of one (SGO 49231) has the altitude 3700 m,
whereas that of SGO 64001 has no elevation,
though it was entered in the JSTOR database as
3880 m. Thellung (1906), Muñoz-Pizarro (1960),
and Boelcke (1964) did not lectotypify the species,
and the more complete sheet is designated above
as the lectotype.
Hitchcock (1945) reduced this native South
American species to synonymy of the Eurasian
weed Lepidium ruderale, and reported the later
from Argentina based on Fischer 37, which is
cited above. Both species have pinnatisect basal
leaves and similar fruit shape and size. However,
L. myrianthum differs by having 1-pinnatisect (vs.
(1-)2-3-pinnatisect) basal leaves, reticulate (vs.
non-reticulate) fruits, stems puberulent with cur-
ved vs. straight papillae, and fruiting pedicels gla-
brous or puberulent only adaxially (vs. puberulent
all around).
Lepidium nitidum Nutt. in Torr. & A. Gray, Fl.
N. Amer. 1: 116. 1838. TYPE: United States.
California, Santa Barbara, T. Nuttall s.n.
(holotype BM!; isotype NY!).
Lepidium chilense Kunze ex Walp., Nov. Act. Nat. Curr.
19, suppl. 1: 250. 1843. Lepidum spicatum var. chi-
lense (Kunze ex Walp.) Reiche, Anales Univ. Chile
90: 93. 1895. TYPE: Chile, sine data, Meyen s.n.
(holotype B!).
Lepidium tenuifolium Phil., Anales Univ. Chile. 81: 333.
1892. Lepidum bipinnatifidum var. tenuifloium
(Phil.) Reiche, Anales Univ. Chile 90: 95. 1895.
TYPE: Chile. Near Chillán, M. A. Solis s.n. (holoty-
pe SGO-63989!).
Lepidium tenuissimum Steud., Nomen. Bot. 2, 28. 1941.
TYPE. Chile. Valparaíso, VII-1830, C. G. L. Bertero
1081 (holotype P!; isotypes BM!, GH!).
Lepidium curicoanum Phil., Anales Univ. Chile 81: 334.
1892. Lepidium bipinnatifidum var. curicoanum
(Phil.) Reiche, Anales Univ. Chile 90: 95. 1895.
TYPE: Chile. Curicó, 1891, M. Vidal s.n. (holotype
SGO-071467!).
Distribution. South America (Chile) and disjunct
in North America (California).
Observations. Hitchcock (1945) recognized
Lepidium curicoanum both as a distinct species
and as a synonym of L. nitidum. Examination of
the types of both species reveals that they are
conspecific.
Lepidium parodii Thell., Repert. Spec. Nov.
Regni Veg. 21: 254. 1925. TYPE: Argentina.
Buenos Aires, Avellaneda, 18-X-1924, L. R.
Parodi 5840 (lectotype Z! here designated;
duplicate BAA!).
Distribution. Endemic to Argentina.
Observations. Thellung (1925) listed two
syntypes in the original description of the species,
and the more complete collection is taken as the
lectotype.
Lepidium pedersenii Al-Shehbaz, sp. nov.
TYPE: Argentina. Chaco, Depto. San Fernan-
do, Isla Soto, low floodable ground, 25-VIII-
1967, T. M. Pedersen 8352 (holotype MO;
isotype BAA). Fig. 3.
Lepidium calycinum Godron var. integrifolium Thell.,
Repert. Spec. Nov. Regni Veg. 11: 309. 1913. L. ale-
tes J. F. Macbr. var. integrifolium (Thell.) Boelcke,
Parodiana 4: 58. 1986; not L. integrifolium Nutt. ex
Torr. & A. Gray, Fl. N. Amer. 1: 116. 1838. TYPE:
Paraguay. Pilcomayo, 1906, T. Rojas 368 (lectotype
B! here designated; duplicate BM!).
Differt a Lepidio auriculato foliis basalibus
serratis vel serrulatis (non 1-3-pinnatifidis vel -
pinnatisectis), foliis caulinis subintegris vel serra-
tis, et pedicellis fructiferibus abaxaliter glabris.
156
DARWINIANA 48(2) 141-167. 2010
157
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
Fig. 2. Lepidium hickenii. A, plant. B, flower. C, fruit. Scales: A = 3 cm, B = 0.5 mm, C = 1 mm. Drawn by the author
from C. M. Hicken s.n. (holotype SI).
Herbs, annual, often strigillose with trichomes
0.2-0.5 mm. Stems 1.5-4.5 dm, erect, usually diva-
ricately branched above, strigillose with retrorse
or rarely spreading trichomes. Basal leaves 2-8(-
11) × 0.3-1.5(-2.5) cm; petioles 0.5-3.5(-5) cm;
leaf blade oblanceolate, glabrous or pubescent
only along midvein, margin serrate to serrulate
along entire length, ciliate; middle cauline leaves
1-2.5 × 0.2-0.6 cm, sessile, base auriculate, margin
subentire to serrate. Racemes dense, retrorse or
rarely spreading strigillose; fruiting pedicels slen-
der, strongly recurved about middle, ascending at
base, divaricate distally, 2-3 mm, narrowly win-
ged, strigillose adaxially and ciliate at margin.
Sepals oblong, 0.5-0.8 mm, persistent, apex white,
puberulent outside; petals white, 0.3-0.5 × 0.03-
0.05 mm; stamens 2, median; filaments 0.5-0.9
mm; anthers 0.1-0.2 mm. Fruits dehiscent, obova-
te-orbicular, 2.5-3.5 × 2-3 mm, winged apically,
glabrous, apex deeply and broadly emarginate;
apical notch 0.4-0.5 mm; style obsolete or to 0.05
mm, included in apical notch. Seeds reddish
brown, ovate-oblong, usually narrowly margined
or winged at least distally, 1.2-1.4 × 0.7-0.8 mm,
margined; cotyledons incumbent.
Etymology. The species is named in honor of
Troels Myndel Pedersen (26 September 1916 - 5
May 2000) who collected the type gathering and
several paratypes cited below.
Distribution and habitat. Lepidium pedersenii
grows on flood plains and along roadsides and
river banks in northeastern Argentina (Chaco,
Corrientes, Formosa, Misiones, Santa Fé, and
Tucumán) and adjacent Paraguay.
Observations. Lepidium pedersenii was treated
by Thellung (1906) and Boelcke (1986) as variety
integrifolium of L. calycinum and L. aletes, res-
pectively. As shown above, the latter two species
are synonymized herein under L. auriculatum.
Both L. pedersenii and L. auriculatum are erect
annuals with persistent sepals and similar fruit
morphology. However, the new species is easily
distinguished by having primarily serrate to serru-
late [vs. (1 or) 2 (or 3)-pinnatifid or -pinnatisect to
pectinate] basal leaf margin, subentire or serrate
(vs. pinnatifid to pinnatisect or pectinate) cauline
leaves, and fruiting pedicels glabrous abaxially
(vs. puberulent all around). In three collections,
Serafín & Pierotti 6545 (GH, LIL), Zardini et al.
860 (LP), and Schwindt 710 (GH, LIL), both spe-
cies are mounted on the same sheet, but in others
(e.g., Montes 1086 vs. Montes 1087 and Schwindt
502 vs. Schwindt 503) the first and second acces-
sion of each collection are L. pedersenii and L.
auriculatum, respectively.
The collection of Lepidium calycinum var. inte-
grifolium designated herein as the lectotype is by
far the most complete of all syntypes cited by its
original author.
Paratypes
ARGENTINA. Chaco. Depto. 1 de Mayo,
Boelcke & Correa 14479 (BAA, BACP); Colonia
Benítez, Schulz 145 (BAB). Corrientes. Barranca
del río Paraná, Rojas 11525 (LIL). Depto. Bella
Vista: Ruta 27, 10 km S of Bella Vista, Toropí,
Schinini & Cristóbal 9839 (G, SI). Depto. Empe-
drado, El Pollo 2 leguas al Este, Ibarrola 3168
(GH, LIL); Estancia La Yela, Pedersen 12975
(BACP). Depto. Itatí, Pedersen 7067 (MO).
Depto. Mburucuyá: Estancia Santa Teresa, Peder-
sen 1813 (LP, MO); Estancia Santa Teresa, Bur-
kart 19329 (BACP, SI, US), Pedersen 2 (MO).
Depto. Santo Tomé, Río Uruguay y Arroyo Chimi-
ray, Schinini & Ahumada 20843 (BAA). Formosa.
Formosa, Jörgensen 2596 (MO); Depto. Laishi,
Ruta Nac. 11, Río Salado, 4 km S Tatané, Boelcke
13342 (BAA), Boelcke 13338 (BAA). Misiones.
San Javier, Aug 1902, Burmeister s.n. (BAB);
Posadas, La Granja, Ekman 1996 (F, P). Depto.
Candelaria: Santa Ana, Montes 1086 (GH, LIL);
Loreto, Montes 14670 (CAS, NY); Loreto, Montes
508 (BAA, BAB). Depto. Cainguás: Mineral,
Schwindt 710 (GH, LIL); Puerto Rico, Schwindt
502 (LIL). Depto. Capital: Garupá, Grondona &
Spegazzini 1310 (BAB). Depto. Bernardo de Irigo-
yen, San Antonio, Serafín & Pierotti 6545 (GH,
LIL). Depto. Iguazú: Victoria, Schwindt 2835
(LIL); Parque Nacional Iguazú, Zardini et al. 852,
860 (LP); Parque Nacional Bañado, Múlgura et al.
577 (BAA). Santa Fe. Depto. Vera, Arroyo
Golondrinas, Cristóbal et al. 2007 (BAA). Tucu-
mán. Depto. Leales: Chañar Pozo, Venturi 420
(LIL). PARAGUAY. Boquerón, Eas. Pozo Once,
Colonia Menno, Vanni et al. 1785 (CTES, F);
158
DARWINIANA 48(2) 141-167. 2010
Capitán Miranda, Lurvey 212 (MO). Depto. A.
Paraná-Viv. Ftal. Itaipú, Itaipú Binacional 24
(MO); Puerto Stroessner, Krapovickas & Cristó-
bal 13392 (BAA).
Lepidium perfoliatum L., Sp. Pl. 2: 643. 1753.
TYPE: "Habitat in Persia, Syria." [Lectotype
Herb. Clifford: 331, Lepidium 3 (BM-
000646274) designated by B. Jonsell & C. Jar-
vis, Nord. J. Bot. 22: 70. 2002].
Distribution. Native to N Africa, Asia, and Euro-
pe; introduced into South America (Argentina),
North America, and Australia.
Lepidium philippianum (Kuntze) Thell.,
Neue Denkschr. Schweiz. Naturf. Ges. 41:
200. 1906. Nasturtium philippianum Kuntze,
Revis. Gen. Pl. 1: 937. 1891. Based on Lepi-
dium suffruticosum Phil., Linnaea 20: 670.
1856, non L., Mant. 1: 91. 1767. TYPE: Chile.
Cordillera de Yerba Loca, R. Philippi s.n.
(holotype SGO-63998!).
Lepidium philippianum var. brachystylum Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 200. 1906.
TYPE: Chile, Cordillera de Santiago, R. Philippi 630
(holotype LE; isotype K!).
Onuris reichei Gilg & Muschl., Bot. Jahrb. Syst. 42:
467. 1909. TYPE: Chile. Cordillera de Santiago,
2700 m, C. Reiche s.n. (holotype B!).
Distribution. Endemic to Chile.
Observations. The limits of Lepidium philippia-
num were broadly delimited by both Thellung
(1906) and Hitchcock (1945) to also include (as
var. boliviense) the Bolivian endemic L. beckii.
Lepidium pinnatifidum Ledeb., Fl. Ross. 1:
206. 1841. TYPE: Russia, "Inter plantas astra-
chanensi," Blume D19 (holotype LE!, plant on
the left).
Distribution. Native to Central Asia and Europe;
naturalized in South America (Argentina) and
North America (California).
Lepidium pseudodidymum Thell. ex Druce,
Bot. Exch. Club Soc. Brit. Isles 3: 388. 1914.
TYPE: [Argentina]. Plants alien to Scotland,
Twedside, Galashields, Sekrikshire, 3-IX-1913,
I. M. Hayward s.n. (holotype E!).
Senebiera australis Hook.f., Fl. Antarct. 2: 241. 1843,
not L. australe Kirk, Trans. N. Z. Inst. 14: 381. 1882.
Coronopus pinnatifidus (DC.) Gaertn. var. australis
(Hook.f.) Reiche, Fl. Chile 1: 67. 1896. Coronopus
australis (Hook.f.) Speg., Ann. Mus. Nac. Hist. Nat.
Bs. Aires 7: 227. 1902. Senebiera pinnatifida DC.
var. australis (Hook.f.) Wildman, Exp. Antarct.
Belge, Voyage S. Y. Belgica, Bot. 95. 1905. Corono-
pus didymus (L.) Sm. subsp. australis (Hook.f.) Gilg
& Muschl., Bot. Jahrb. Syst. 42: 449. 1909. TYPE:
Chile. Chonos Archipiélago, XII-1834, C. Darwin
34.bis (holotype K!).
Lepidium inclusum O. E. Schulz, Repert. Sp. Nov.
Regni Veg. 33: 189. 1933. TYPE: Argentina. Tierra
del Fuego, Río Grande, 12-I-1933, A. Castellanos
7757 (holotype B!).
Distribution. Patagonian Argentina and Chile.
Lepidium pubescens Desv., J. Bot. Agric. 3:
180. 1915. TYPE: Peru. Para, Dombey s.n.
(holotype P!).
Lepidium rainmondii O. E. Schulz, Notizbl. Bot. Gart.
Berlin-Dahlem 10: 727. 1929, syn. nov. TYPE: Peru.
Cajamarca, Prov. Contumazá, Cascas, Cerro de Cata-
che, 5500 ft [ca. 1800 m], 25-V-1875, A. Raimondi
7538 (holotype B!).
Lepidium demissum C. L. Hitchc., Lilloa 11: 121. 1945,
syn. nov. TYPE: Bolivia. La Paz, 3800 m, 3-IV-1919,
O. Buchtien 4475 (holotype GH!; isotypes US!).
Lepidium scabrifructum C. L. Hitchc., Lilloa 11: 119.
1945, syn. nov. TYPE: Bolivia. Atocha, 20-III-1921,
3700 m, E. Asplund 6205 (holotype US!; isotype S!).
Distribution. Bolivia and Peru.
Observations. Neither Schulz (1929) nor Hitch-
cock (1945) examined the type of Lepidium pu-
bescens. The types of L. raimondii, L. demissum,
and L. scabrifructum are indistinguishable from
that of L. pubescens in every aspect of foliage,
flowers and fruits. The only difference is the pre-
sence in L. demissum of nearly persistent sepals
and in L. scabrifructum only slightly smaller (ca.
159
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
3.5 × 3 mm vs. 3.5-5 × 3-4.5 mm) fruits. The pre-
sence of minute trichomes on the margin of fruit
valve, annual habit, and pinnatifid or pinnatisect
leaves should readily distinguish L. pubescens
(including its new three synonyms) from the other
congeners.
Lepidium quitense Turcz., Bull. Soc. Imp.
Naturalistes Moscou 27(2): 39. 1854. TYPE:
Ecuador. Quito, Plains of Pomasqui, 1850,
Jameson 892 (holotype KW; isotypes G-BOIS!,
G-DEL!, US!; fragment F!).
Lepidium quitense var. integrifolium Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 213. 1906.
TYPE: Ecuador. Guallabamba, XII-1886, A. Sodiro
56 (holotype B!).
Lepidium quitense var. microphyllum Thell., Neue
Denkschr. Schweiz. Naturf. Ges. 41: 213. 1906.
TYPE: Ecuador. Sine data, A. J. A. Bonpland s.n.
(holotype B-W!; isotypes B!, P!).
Distribution. Endemic to Ecuador.
Lepidium rahmeri Phil., Anales Mus. Nac.
Chile 1891: 5. 1891. TYPE: Chile. [Región I],
Tarapacá, Calcalhuai, C. Rahmer s.n. (holotype
SGO-63996!; isotype SGO-71461!).
Distribution. Argentina (new), Chile.
Observations. The species is reported for the
first time from Argentina based on the many
collections cited below, all of which were pre-
viously undetermined. The species resembles L.
argentinum in having acute fruit valves and often
minutely puberulent valve keel. However, it dif-
fers in the perennial (vs. annual) habit and in
having orbicular to ovate-orbicular (vs. oblong to
oblong-ovate) fruits.
Specimens examined
ARGENTINA. Jujuy. San Antonio de los
Pibes, Sleumer 3241 (BAA, LIL, SI); Huacalera,
Cabrera 12066 (LP). Depto. Cochinoca, Cerro
Huauca, Charpin & Novara 23129 (G). Depto.
Humahuaca, Azul Pampa, Cabrera et al. 21411
(LP, MO); Humahuaca, Parodi 9668 (BAA);
Cuesta de Azul Pampa, J. H. Hunziker et al. 10465
(SI); Mina Aguilar, Cabrera et al. 15439 (LP), J.
H. Hunziker et al. 10482 (SI); de Pucará a Palca de
Aparzo, Kiesling 3512, 3633 (SI); Tres Cruces,
Cabrera et al. 27455 (SI). Depto. Santa Bárbara,
Santa Bárbara, Co. Pereyra, Gualianone et al.
1915 (SI). Depto. Santa Catalina, Santa Catalina,
Morrone et al. 2678 (SI). Depto. Tilcara, Quebra-
da de Huasamayo, Cabrera et al. 31571 (SI, US).
Depto. Tumbaya, Quebrada de Lipán, Zuloaga &
Deginani 3573 (SI); camino de Purmamarca al
Abra de Lipán, R. Kiesling et al. 5192 (SI), Kiel-
sing et al. 5221 (SI); 32 km de Purmamarca,
Morrone et al. 2730 (SI); Abra de Lipán, Cabrera
et al. 31674 (SI), Cabrera et al. 27408 (SI); Vol-
cán, Chilcayo, camino a Abra Morada, Kiesling et
al. 5819 (SI); Ciénaga Grande, 11 km S El More-
no, Nicora et al. 8911 (SI); camino de Purmamar-
ca a Abra de Lipán, Abra Blanca, Kiesling et al.
5303 (SI), Kiesling 5310 (SI); subida de Purma-
marca a Abra de Pives, Cabrera et al. 26356 (SI);
El Moreno, Angosto del Chañi, ca. Incahwasi,
Kielsing et al. 5255 (SI). Depto. Yavi, La Quiaca,
Novara 8259 (G); Pumahuasi, Giusti et al. MC551
(BAA). Salta. Puerto Tastil, Feb 1943, Pedraluer-
ca s.n. (BAA). Depto. Cachi, ruta 33, km 44 des-
pués de Cachi, Charpin & Lazare 23995 (G).
Lepidium reichei Phil. ex Reiche, Fl. Chile 1:
64. 1896. TYPE: Chile. Andes de Santiago,
Valle Largo, II-1892, C. Reiche s.n. (holotype
SGO-63994!; isotype B!).
Lepidium brevicaule Barn. in Gay, Hist. Chile Bot. 1: 165.
1845, syn. nov. Non Lepidium brevicaule Hoppe ex W.
D. J. Koch, Deutschl. Fl., ed. 3, 4: 519. 1833. Nastur-
tium brevicaule Kuntze, Revis. Gen. Pl. 2: 937. 1891.
Lepidium barneoudianum Sukorý, Gayana Bot. 60(2):
135. 2003. TYPE: Chile. Cordilleras de Coquimbo,
3300 m, 1839, C. Gay 335 (holotype P!).
Lepidium morrisonii C. L. Hitchc., Lilloa 11: 116. 1945,
syn. nov. TYPE: Chile. Coquimbo, Río Ojotas, NE of
La Vega Redonda, Andes back of Cuncumen, 3000
m, 25-II-1939, J. L. Morrison & R. Wagenknecht
17418 (holotype UC!; isotypes F!, LIL!, G!, MO!,
NA!, SI!).
Lepidium bonariense L. var. gayi Thell., Bull. Herb.
Boissier ser. 2, 8: 914. 1908, syn. nov. TYPE: Chile.
Prov. Coquimbo, 1838, C. Gay s.n. (holotype P!).
160
DARWINIANA 48(2) 141-167. 2010
161
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
Fig. 3. Lepidium pedersenii. A, plant. B, flower. C, fruit. Scales: A = 2 cm, B = 0.5 mm, C = 1 mm. Drawn by the
author from T. M. Pedersen 8352 (holotype MO).
Distribution. Endemic to Chile.
Observations. A very distinctive species easily
distinguished by its small fruits, straight fruiting
pedicels subappressed to rachis, perennial habit,
and pinnatisect leaves. Hitchcock (1945) suspec-
ted that Lepidium morrisonii might be conspecific
with L. brevicaule, but he refrained from treating
them as such because he did not examine the type
of the latter.
Teillier (1993) has correctly demonstrated that
Lepidium reichei is rather distinct morphologically
from L. philippianum, a species under which
Hitchcock (1945) erroneously treated L. reichei as
a synonym. However, Teillier (1993) failed to
synonymize L. morrisonii and the later homonym
L. brevicaule and its new name L. bareoudianum
(Sukorý, 2003) with L. reichei.
Lepidium rhytidocarpum (Hook.) Al-Shehbaz,
Novon 12: 9. 2002. Senebiera rhytidocarpa
Hook., London J. Bot. 2: 506. 1843. Coronopus
rhytidocarpus (Hook.) Macloskie, Rep. Princ.
Univ. Exped. Patagonia, Botany 8: 428. 1905.
TYPE: Argentina. "Patagonia", Tweede s.n.
(holotype K!).
Distribution. Argentina and Uruguay.
Lepidium santacruzensis Al-Shehbaz, sp. nov.
TYPE: Argentina. Santa Cruz, Depto. Deseado,
Puerto Deseado, Isla Quirogua, Desembocadu-
ra de la Ría, 17-XI-1963, M. N. Correa, L. Men-
donza & C. Movia 2541 (holotype BAA; isoty-
pe BAB). Fig. 4.
Differt a Lepidio auriculato et L. pedersenii
caulibus prostratis, petalis nullis, fructibus 2.5-2.7
(vs. 3-3.7) mm longis, apice emarginatibus stylo
equilongis, et seminibus non-alatis nec margina-
tis.
Herbs, annual, hirsutulous on stems, pedicels
and sepals with straight trichomes 0.1-0.4 mm.
Stems 7-15 cm, prostrate, branched an base and
throughout. Basal leaves soon withered, oblanceo-
late, 1-2.5 cm, narrowed to petiolar base, entire or
few toothed; cauline leaves sessile, 7-15 × 3-7
mm, oblong, coarsely and obtusely dentate,
strongly auriculate at base, glabrous. Racemes
dense, slightly elongated in fruit; fruiting pedicels
strongly recurved about middle, erect at base, flat-
tened, divaricate distally, 2-2.5 mm, puberulent all
around, narrowly winged. Sepals ovate, 0.6-0.7
mm, persistent, apex white, pubescent outside;
petals absent; stamens 2, median; filaments 0.6-
0.8 mm; anthers ca. 0.15-0.2 mm. Fruits dehiscent,
obovate-orbicular, 2.5-2.7 × 2.2-2.5 mm, narrowly
winged apically, glabrous, apex emarginate; apical
notch ca. 0.1 mm; style ca. 0.1 mm, as long as api-
cal notch. Seeds reddish brown, ovate-oblong,
wingless, not margined, 1.1-1.2 × ca. 0.7 mm;
cotyledons incumbent.
Etymology. The species is named after Santa
Cruz Province.
Distribution and habitat. Known only from
the type collection.
Observations. The type material of Lepidium
santacruzensis was cited by Boelcke & Romanc-
zuk (1984) as L. aletes (herein as L. auriculatum).
However, it resembles that species only in having
auriculate cauline leaves, persistent sepals, and
fruiting pedicels puberulent all around. From L.
auriculatum and the related L. pedersenii, L. san-
tacruzensis is readily distinguished by the lack (vs.
presence) of petals and by having prostrate (vs.
erect) stems branched at the base and throughout
(vs. branched only distally), caducous (vs. persis-
tent) basal leaves, smaller fruits 2.5-2.7 (vs. 3-3.7)
mm, apical notch ca. 0.1 mm (vs. 0.3-0.5 mm),
style as long as the apical notch (vs. obsolete and
included in apical notch), and wingless and not
margined (vs. at least apically winged or margi-
ned) seeds. Furthermore, it differs from L. auricu-
latum by having entire or toothed vs. (1 or) 2 (or
3)-pinnatifid or -pinnatisect] leaves. From L.
pedersenii, the new species differs by the lack (vs.
presence) of distinct petiole on basal leaves and
entire or few-toothed (vs. serrate or serrulate along
entire margin).
Lepidium sativum L., Sp. Pl. 2: 644. 1753.
TYPE: [lectotype Herb. Linn. 824.11 (LINN)
162
DARWINIANA 48(2) 141-167. 2010
designated by W. Fawcett & A. B. Rendle, Flora
of Jamaica 3: 243. 1914].
Distribution. Native to N Africa and Eurasia;
cultivated and naturalized in South America
(Argentina), North America, and Australia.
Lepidium serratum (Poir.) Al-Shehbaz, Novon
12: 9. 2002. Senebiera serrata Poir., Encycl.
(Lamarck) 7: 76. 1806. Coronopus serratus
(Poir.) Desv., J. Bot. Agric. 3: 163. 1815. TYPE:
Uruguay. Montevideo, Commerson s.n. (holotype
P-JU; isotypes fragments BAA!, P[3]!).
Distribution. Argentina and Uruguay.
Lepidium solomonii Al-Shehbaz, Ann. Missou-
ri Bot. Gard. 73: 830. 1986. TYPE: Bolivia. La
Paz, Prov. Los Andes, 6.6 km NW of Batallas
on the principal road along Lake Titicaca,
16º15’S, 68º33’W, 3,850 m, rocky hillside, 5-
II-1984, J. C. Solomon 11448 (holotype MO!;
isotypes GH!, LPB).
Distribution. Endemic to Bolivia.
Lepidium spathulatum Phil., Fl. Atacam. 8. 1860.
Nasturtium spathulatum (Phil.) Kuntze, Revis.
Gen. Pl. 1: 937. 1891. TYPE: Chile. Cachinal de la
Costa, R. Philippi s.n. (holotype SGO-063990!).
Distribution. Endemic to Chile.
Lepidium spicatum Desv., J. Bot. Agric. 3: 178.
1815. TYPE: Magallanes, Commerson s.n.
(holotype P!; isotype P!). The more complete
specimen annotated by Desvaux's handwriting
is taken herein as the type.
163
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
Fig. 4. Lepidium santacruzensis. A, plant. B, fruit. Scales: A = 1 cm, B = 1 mm. Drawn by the author from M. C.
Correa et al. 2541 (holotype BAA).
Lepidium racemosum Griseb., Abh. Königl. Ges. Wiss.
Göttingen 6: 116. 1854. Lepidium spicatum var.
racemosum (Griseb.) Boelcke, Fl. Patag. 8(4a): 468.
1984. TYPE: Peninsula Brunswick, Oazy Harbor, W.
Lechler 1114 (lectotype GOET! designated by C. L.
Hitchcock, Lilloa 11: 101. 1945; duplicates G!,
P[2]!).
Lepidium spicatum var. calyx-persistente Boelcke, Paro-
diana 3: 26. 1984, syn. nov. TYPE: Argentina. Entre
Ríos. Depto. Gualeguaychú, Pto. Constanza, arroyo
Carquejas, 9-IV-1960, A. Burkart & J. C. Gamerro
21749 (holotype SI!; isotype BAA!).
Distribution. Argentina and Chilean Patagonia.
Observations. Both Thellung (1906) and Hitch-
cock (1945) reduced Lepidium racemosum to
synonymy of L. spicatum, but Boelcke & Romanc-
zuk (1984) treated it as a variety of the latter. The
main differences they used to separate the two
varieties is the presence in var. racemosum of
minute auricles on the upper pinnately divided
cauline leaves (vs. non-auriculate and entire or tri-
fid upper leaves). However, this variation is conti-
nuous, and mixed plants of both "varieties" are
found in many collections.
Lepidium steinbachii O. E. Schulz, Notizbl.
Bot. Gart. Berlin-Dahlem 9: 1037. 1926. TYPE:
Bolivia. Santa Cruz, Prov. Cercado, Pampitas
del Río Piray, 450 m, 24V-VIII-916, J. Stein-
bach 2724 (holotype B!; isotypes GH!, LIL!,
SI!).
Distribution. Endemic to Chile and known only
from the type collection above.
Lepidium strictum (S. Wats.) Rattan ex B. L.
Rob. in A. Gray, Syn. Fl. N. Amer. 1(1): 129.
1895. Lepidium oxycarpum Torr. & A. Gray var.
strictum S. Wat. in Brewer & S. Watson, Bot.
California 1: 46. 1876. TYPE: United States.
California. El Dorado County, near Placerville,
1878, V. Rattan s.n. (holotype GH!).
Distribution. Apparently native to Chile and per-
haps Peru, also in the western United States.
The disjunct distribution of Lepidium strictum
in both the United States (California, Oregon) and
Chile requires studies to show if the species is
indeed native to South America as suspected by
Hitchcock (1945).
Lepidium stuckertianum (Thell.) Boelcke,
Parodiana 4(1): 40. 1986. Lepidium bonariense
L. var. stuckertianum Thell., Repert. Spec. Nov.
Regni Veg. 13: 302. 1914. TYPE: Argentina.
Santiago del Estero, Salavina, T. Stuckert 7434
(lectotype Z designated by O. Boelcke, Paro-
diana 4: 40. 1986; duplicate CORD!).
Distribution. Endemic to Argentina.
Observations. Thellung (1914) erroneously lis-
ted the above lectotype as Stuckert 7439 instead of
7434.
Lepidium tandilense Boelcke, Darwiniana
13: 521. 1964. TYPE: Argentina. Buenos Aires,
Tandil, Sierras de las Ánimas, 14-X-1962, O.
Boelcke, M. N. Correa, N. M. Bacigalupo 2441
(holotype BAA!; isotypes BAB!, SI!).
Distribution. Endemic to Argentina.
Lepidium trianae Thell., Neue Denkschr.
Schweiz. Naturf. Ges. 41: 214. 1906. TYPE:
Colombia. Nouvelle Grenada [Cundinamarca],
Bogota, 1851-1857, 2700 m, J. Triana s.n. (lec-
totype G-DC! designated by C. L. Hitchcock,
Lilloa 11: 98. 1945 and herein; duplicates BM!,
G-DC!, K!, NY!, P!).
Distribution. Endemic to Colombia.
Observations. Hitchcock did not examine the lec-
totype he designated, and the more complete plant
with flowers and fruits and annotated in Thellung's
handwriting as "Lepidium trianae Thell. n.sp." is
confirmed herein as the lectotype. The G-DC dupli-
cate without flowers and fruits is taken as the isolec-
totype. Thellung (1906) cited one collection (Wed-
dell s.n.; P) from Bolivia, and although I have not
examined that collection, it is highly unlikely that
the species extends its range that far south.
164
DARWINIANA 48(2) 141-167. 2010
Lepidium virginicum L., Sp. Pl. 2: 645. 1753.
TYPE: "Habitat in Virginia, Jamaicae glareo-
sis" [lectotype Herb. Linn. # 824.18 (LINN)
designated by W. Marais, Flora of Southern
Africa 13: 94. 1970].
Lepidium danielsii C. L. Hitchc., Lilloa 11: 123. 1945,
syn. nov. TYPE: Colombia. Depto. Antioquia. Mede-
llín, VIII-1933, H. Daniels s.n. (holotype NY!).
Distribution. Native to North America, introdu-
ced into South America (Bolivia, Brazil, Colom-
bia, Ecuador, Paraguay, Peru, Surinam, Venezue-
la), Europe, Asia, and Australia.
Observations. Lepidium danielsii is indistinguis-
hable from the highly variable L. virginicum in
every aspect of the plant. However, its type has
somewhat large fruits, a feature that occurs spora-
dically throughout the native and naturalized ran-
ges of L. virginicum.
Lepidium werffii Al-Shehbaz, sp. nov. TYPE:
Peru. Arequipa-Ubinas road, 16°23'13" S,
71°20'47" W, 2600-3400 m, scrub, 17-IV-2006,
H. van der Werff, L. Valenzuela, & E. Sculli
20743 (holotype MO; isotypes B, CAS, F, GH,
K, SI, P, US). Fig. 5.
Differt a Lepidio meyenii staminibus 4 (vs. 2),
petalis obovatis 1.2-2.2 (vs. 0.3-0.5) mm latis, val-
vis reticulatis, et fructibus 4-5.5 [vs. 2.5-3.8(-4)]
mm latis.
Herbs, perennial, sparsely to densely puberu-
lent with straight, spreading trichomes 0.01-0.2
mm; roots not fleshy; caudex simple, 0.5-2 cm in
diam. Stems 4-25 cm, decumbent to ascending,
often several from caudex, branched above. Basal
leaves 1.7-11 cm; petioles 1-6 cm, persistent, stra-
mineus, flattened at base; blade undivided and
oblanceolate with 5-7 subapical teeth, or pinnatifid
and with 2-4 dentate to incised lateral lobes on
165
I. A. AL-SHEHBAZ. A synopsis of the South American Lepidium (Brassicaceae)
Fig. 5. Lepidium werffii. A, plant. B, basal leaf. C, flower (two petals removed). D, fruit. Scales: A, B = 2 cm, C, D =
1 mm. Drawn by the author from H. van der Werff et al. 20743 (holotype MO).
each side; cauline leaves shortly petiolate to sub-
sessile, laciniate, not auriculate at base. Racemes
lax, terminal, elongated considerably in fruit;
rachis puberulent with straight trichomes; fruiting
pedicels slender, straight, spreading, 4-7 mm,
narrowly winged, glabrous abaxially. Sepals
oblong, 1.5-2 mm, persistent; petals white, obova-
te, 2-3.5 × 1.2-2.2 mm, longer than sepals; sta-
mens 4, median; filaments 1-1.5 mm; anthers 0.4-
0.5 mm. Fruits dehiscent, broadly rhombic, 2.5-4.5
× 4-5.5 mm, distinctly wider than long, subcons-
tricted below apex, glabrous, distinctly veined,
narrowly winged apically, apex emarginate; apical
notch 0.05-0.2 mm; style 0.5-1 mm, exserted from
apical notch. Seeds brown, ovate-oblong, win-
gless, 1.4-1.8 × 0.8-1 mm; cotyledons incumbent.
Etymology. Lepidium werffii is named in honor
of Henk van der Werff (1946- ), the world expert
on the family Lauraceae and one of the collectors
of the type gathering.
Distribution and habitat. The species grows in
scrub and puna vegetation at 2600-4200 m in sou-
thern Peru.
Observations. Lepidium werffii is most closely
related to L. meyenii, from which it differs by
having obovate (vs. linear to oblanceolate) petals
1.2-2.2 (vs. 0.3-0.5) mm wide, four (vs. two) sta-
mens, broadly rhombic (vs. elliptic to rhombic-
suborbicular) fruits 4-5.5 [vs. 2.5-3.8(-4)] mm
wide and distinctly wider (vs. narrower) than long,
and veined (vs. not veined) fruit valves.
Paratypes
PERU. Arequipa. Arequipa-Ubinas road,
Werff, et al. 20812 (BM, E, G, MO, NY). Tacna.
Prov. Tarata, Cord. del Barroso, Torre et al. 2111
(MO).
ACKNOWLEDGEMENTS
I thank Drs. Ana Anton (CORD), Vladimir Dorofe-
yev (LE), Martin Gardner (E), Dmitry German (ALTB),
Jochen Heinrich (GOET), Sergei Mosyakin (KW),
Melica Muñoz-Schick (SGO), Reto Nyffeler (Z), Joana
Osborne (K), Rusty Russell (US), Robert Vogt (B),
Emily Wood and Walter Kitteredge (GH), Thomas A.
Zanoni (NY), and Hans-Joachim Zündorf (JE) for their
help in locating type collections in their herbaria. I am
also grateful to all the curators and directors of herbaria
cited above for the loan of specimens. I also thank Fred
Keusenkothen for helping with the digital imaging of
the new species. Fieldwork and herbarium studies in
Argentina were generously supported by the National
Geographic Society (Grant 8139-06). Last but certainly
not least, I thank Fernando O. Zuloaga and Raúl E. Poz-
ner for the advice on the manuscript.
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... With ca 265 species (German 2020), Lepidium is the second-largest and one of just three naturally cosmopolitan genera of the family (Al-Shehbaz 2017). As currently delimited (Al-Shehbaz 2010, 2012a, b, 2015, Lepidium includes not only species with angustiseptate (strongly compressed at right angle to the septum) and dehiscent fruits (i.e. Lepidium in a traditional sense), but also representatives with didymous, quadrangular, terete or inflated, sometimes indehiscent pods, but even with such a broad concept, fully supported phylogenetically (Mummenhoff et al. 2001(Mummenhoff et al. , 2009, it is distinguished from other mustard genera by a combination of fruit silicles with one (very rarely two) subapical ovules per locule, often copiously mucilaginous seeds with almost exceptionally incumbent cotyledons, development of usually both lateral and median nectar glands and indumentum of no or only simple trichomes Gaskin 2010, Al-Shehbaz 2015). ...
... Lepidium in a traditional sense), but also representatives with didymous, quadrangular, terete or inflated, sometimes indehiscent pods, but even with such a broad concept, fully supported phylogenetically (Mummenhoff et al. 2001(Mummenhoff et al. , 2009, it is distinguished from other mustard genera by a combination of fruit silicles with one (very rarely two) subapical ovules per locule, often copiously mucilaginous seeds with almost exceptionally incumbent cotyledons, development of usually both lateral and median nectar glands and indumentum of no or only simple trichomes Gaskin 2010, Al-Shehbaz 2015). The genus has centers of diversity and endemism on all continents except for Antarctica (Mummenhoff et al. 2004, Al-Shehbaz 2010; within Eurasia, southwest Asia is one of these regions (German 2014). ...
... The plants were photographed, collected and crosschecked with treatments covering Iran and neighboring areas (Boissier 1867, Busch 1939, Grossheim 1950, Hedge 1965, 1968, Zohary 1966, Maire 1967, Mouterde 1970, Jafri 1973, Täckholm 1974, Vinogradova 1974, Meikle 1977, Hedge and Lamond 1980, Vasconcellos 1993, Miller 1996, Boulos 1999, Zhou et al. 2001, Snogerup 2002, Dorofeyev 2008, 2012, Bona 2014, Al-Shehbaz 2015, Fakhr Ranjberi 2017, POWO 2020. Having in mind the ability of many Lepidium species, especially annual, to become established in new regions as weedy and ruderal from human-mediated long-distance dispersals (Snogerup 2002), some basic extracontinental accounts such as Marais (1970), Jonsell (1975Jonsell ( , 2000, Hewson (1982), Rollins (1993), Al-Shehbaz (2010, 2012b, Al-Shehbaz and Gaskin (2010) and de Lange et al. (2013) were also consulted. Besides, data available in the virtual herbaria B, E, G, P and W via relevant databases Curators Herbarium B (2000+), CGH (2020), MNHN (2020), RBGE (2020) and Virtual Herbaria (2020), were considered. ...
Article
Lepidium khalkhalicum, a new species from northwestern Iran, is described. It appears to be a local endemic restricted to open slopes of marl hills in the Khalkhal region (Ardabil province). The novel species is easily distinguished from similar species by a set of both vegetative and generative characters such as less developed indumentum, undivided leaves, much longer style, more divaricate, non‐thickened pedicels exceeding silicles, poorly developed fruit wing, etc. A detailed description accompanied by color photographs, distribution and habitat data, and a proposed conservation status of the new species are provided. The particular interest of the discovery in view of its close relationship to the garden cress, L. sativum, is stressed.
... En América del Sur, se reportan 50 especies nativas y 12 naturalizadas. En Chile se encuentran 18 especies (Al-Shehbaz, 2010). ...
... Fruto, una cápsula de forma oval. Semillas numerosas, planas, ovales, de 1 mm de largo, y de color café rojizos, más oscuro hacia los márgenes, los que son ligeramente rugosos y donde se ubica un apéndice blanquecino (Al-Shehbaz, 2010;Tay et al., 2011). ...
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The southern distribution range in Chile of Lepidium latifolium L. (Brassicaceae) is expande to the Maule Region. The species was previously known only for the regions of Antofagasta and Coquimbo. L. latifolium is considered a highly aggressive weed in the colonization of productive areas.
... Lepidium ruderale (Fig. 64) Lepidium ruderale is found throughout almost all of Europe with the exception of the arctic region, and also occurs in large parts of temperate and subtropical Asia (Thellung 1906, Meusel et al. 1965. It is likely to have been introduced into many places in the world, but due to confusion with similar species, its distribution cannot be outlined with certainty (Jonsell 1975, Hewson 1982, Al-Shehbaz 2010. It is reliably recorded as introduced into North America (Al-Shehbaz & Gaskin 2010). ...
... Lepidium sativum (Fig. 65) Lepidium sativum is native to north-eastern Africa and south-eastern Asia (Thellung 1906). It is cultivated and has become naturalized in Europe, Asia, North America, South America (Argentina) and Australia (Cheo et al. 2001, Al-Shehbaz 2010. In the Czech Republic it is often cultivated but usually only as a vegetable in pots, with seedlings or young plants harvested. ...
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The catalog describes 1362 species of ferns and flowering plants for the department of Arequipa, more than 200 species more as reported in the last systematic inventory by Quipuscoa, Dillon, & Ortíz in 2006. Ninety-five species are mentioned for the first time for Arequipa and 26 species mentioned in the literature for the department were excluded. In addition to a brief description of the species, the catalog includes information on their ecology, distribution, and human use. In addition to information on systematics and phylogeny, the most important synonyms are listed in the appendix.
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This thesis argues for the need for a more comprehensive discussion of biodiversity use in relation to enhancing benefits of this use for biodiverse countries and promoting more equitable sharing of these benefits. The findings from this doctoral research reveal that biodiversity-based innovation is a social shaping process that has resulted in large benefits. The cumulative capability to use species from biodiversity gives meanings that contribute to the species shaping process, with organisations and institutional changes providing direction and increasing the rate of the shaping process. In showing how innovation takes place and how the appropriation of benefits occurs, this research contributes to studies on science policy and innovation in relation, especially, to biodiversity-based innovation.
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La presente contribución constituye un aporte al conocimiento de la flora vascular de Chile, reuniendo en un solo texto la información acerca del significado del nombre de los géneros chilenos de plantas vasculares. Para ello se ha actualizado la lista de géneros nativos válidos a la fecha y se ha recopilado la información eminentemente dispersa en relación con el origen de los nombres. Adicionalmente se incluye información relevante acerca de la distribución de cada género y el número de especies nativas presentes en Chile.This paper is a contribution to the knowledge of the vascular flora of Chile, bringing together in one text the information about the meaning of names of the Chilean genera of vascular plants. For this purpose, we updated the list of valid genera native to Chile and revised all the available and currently dispersed literature in relation to the origin of the names. We also included relevant information about the distribution of each genus and the number of native species present in Chile.
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Este estudio presenta la descripción taxonómica de un ejemplar tipo de la MACA obtenido el 9 de septiembre de 1989 cerca de Huarancaca, ciudad de Cerro de Pasco, del Departamento de Pasco, como Lepidium peruvianum Chacon sp. nov., especie nueva para el género Lepidium (CRUCIFERA), las características que la distinguen de las especies afines especialmente de Lepidium meyenii Walp. y de Lepidium gelidium Wedd. están especificadas en este estudio. Igualmente presenta el estudio fitoquímico de la MACA realizado en 1960-1962, su hábitat y aclimatación a niveles de la costa. Esta especie cuyo nombre es MACA es oriunda de los Andes Centrales del Perú y se la conoce tanto por su valor alimenticio como farmacológico desde antes de la época del incanato. Crece en los Departamentos de Pasco y Junín en altitudes de 4100 y 4300 m sobre el nivel del mar. El clima en esta región es frío y seco durante el año caracterizándose el verano (enero, febrero y marzo) como lluvioso con un promedio de 250,8 mm y con una temperatura ambiental promedio de 4.8 °C. En invierno (junio, julio y agosto) las lluvias son escasas llegando a 34.4 mm y el promedio de la temperatura ambiental es alrededor de 3.5 °C. Los análisis químicos de la raíz realizados en 1960 y 1961 han demostrado la presencia de glúcidos y cuatro tipos de alcaloides, presentadas en cromatografía de papel Whatman. Estos alcaloides que he denominado "Extracto alcaloideo/marzo 1961", son los principios activos que intervienen en la frecuencia de procreación y dela clara y marcada estimulación de la maduración de los folículos de Graaf realizados en experimentos con ratas y en los machos, un aumento en la cantidad de espermatozoides en los tubos seminíferos y aumento de mitosis y espermatogonia. Estos ensayos los he realizado en el Instituto de Patología de la Facultad de Medicina Humana [de la UNMSM] durante los años 1961 y 1962. Las observaciones histológicas e histoquímicas realizado en 1989 y 1990 en el ejemplar tipo han demostrado que esta especie presenta alcaloides y glúcidos, principios químicos que caracterizaron a los ejemplares de San Juan de Jarpa y Carhuamayo del Departamento de Junín en 1960-1962. Igualmente tiene elevada concentración de Ca, P y Fe, siendo éste último junto con los alcaloides el potencial medicinal de la planta. Se incluyen 10 cuadros, 12 diagramas, 94 fotos y 4 protocolos de autopsia.
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In October 1985 we had the exciting opportunity of seeing the South American species in Chile, in their native environment. Over a hectic fortnight's collecting, travelling more than 2,000 kilometres by jeep, we visited over fifty localities between Puerto Montt and Santiago (Chile). We found Cyttaria specimens at about twenty of the sites visited - they were often abundant - and photographed them in their natural habitat, bringing specimens back to compare with those of Darwin and others. Our specimens now reside in the herbarium of the CAB International Mycological Institute (Herb. IMI) in Kew. In all, we observed four species and these are illustrated here from our field photographs.
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The genus Stubendorffia (Brassicaceae) is distinguished from Lepidium solely by the dehiscent vs. indehiscent angustiseptate fruits. By contrast, Winklera is separated from Lepidium by a combination of perennial habit, pinnatisect leaves, yellow flowers, and wingless fruits, characters all of which occur individually and in various combinations within Lepidium. Extensive molecular studies strongly show that Winklera and polyphyletic Stubendorffia are nested within the earlier-published Lepidium and, therefore, the three genera are herein formally united. The new name Lepidium pavlovii and 10 new combinations, L. afghanicum, L. apterum, L. botschantzevii, L. curvinervium, L. lipskyi, L. olgae, L. orientalis, L. patrinoides, L. pterocarpum, and L. silaifolium, are proposed. Lepidium apterum is lectotypified. A complete generic synonymy of Lepidium and an expanded generic description are presented.
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